A revision of European species of Leccinum gray and notes on extralimital species

Abstract

This paper deals with the generic delimitation of Leccinum and the taxonomic and nomenclatural implications of the phylogenetic results presented in previous papers by Den Bakker et al. (2004a, 2004b). 28S nrDNA data, used in the past to answer questions concerning genus delimitation of Leccinum, are re-analysed. The phylogenetic inferences based on 28S nrDNA largely coincide with Leccinum sensu Singer. The only species that is excluded from Leccinum is L. eximium. At least one truffle-like genus, Chamonixia, should be included in Leccinum. Morphological characters proposed in the past to delimit the genus are discussed in the light of the phylogenetic results. It is concluded that there is no single diagnostic morphological character that unites species of the genus Leccinum and that a better sampling, and other genes that provide a finer phylogenetic resolution, are needed to reach a final answer about the genus delimitation of Leccinum and other genera in the Boletaceae. The second part of this paper discusses characters used in Leccinum-systematics in the past and contains a revision of the European species of Leccinum with notes on related North American species. Sixteen species are accepted for the European continent, of which one, Leccinum albostipitatum, is described as new to science.

Full text

PERSOONIA
Volume
18,
Part
4,511-587
(2005)
A
revision
of
European
species
of
Leccinum
Gray
and
notes
on
extralimital
species
Henk+C.
den
Bakker
.&
Machiel+E.
Noordeloos
Nationaal
Herbarium
Nederland,
Universiteit
Leiden
branch,
P.
O.
Box
9514,
2300
RA
Leiden,
The
Netherlands;
bakker@nhn.leidenuniv.nl
This
paper
deals
with
the
generic
delimitation
of
Leccinum
and
the
taxonomic
and
nomenclatural
implications
of
the
phylogenetic
results
presented
in
previ-
ous
papers
by
Den
Bakker
et
al.
(2004a,2004b).
28S
nrDNA
data,
used
in
the
past
to
answer
questions
concerning
genus
delimitation
of
Leccinum,
are
re-
analysed.
The
phylogenetic
inferences
based
on
28S
nrDNA
largely
coincide
with
Leccinum
sensu
Singer.
The
only
species
that
is
excluded
from
Leccinum
is
L.
eximium.
At
least
one
truffle-like
genus,
Chamonixia,
should
be
included
in
Lec-
cinum.
Morphological
characters
proposed
in
the
past
to
delimit
the
genus
are
dis-
cussed
in
the
light
of
the
phylogenetic
results.
It
is
concluded
that
there is
no
single
diagnostic
morphological
character
that
unites
species
of
the
genus
Leccinum
and
that
a
better
sampling,
and
other
genes
that
provide
a
finer
phylogenetic
resolution,
are
needed
to
reach
a
final
answer
about
the
genus
delimitation
of
Leccinum
and
other
genera
in
the
Boletaceae.
The
second
part
of
this
paper
discusses
characters
used
in
Leccinum-
systematics
in
the
past
and
contains
a
revision
of
the
European
species
of
Leccinum
with
notes
on
related
North
American
species.
Sixteen
species
are
accepted
for
the
European
continent,
of
which
one,
Leccinum
albostipitatum,
is
described
as
new
to
science.
1)
Boletus
corsicus
Roll,
in
Bull.
Soc.
Mycol.
France
12
(1896)
1;
Leccinum
corsicum
(Roll.)
Singer,
Die
Rohrlinge
2
(1967)
87.
2)
Boletus
lepidus
Bouchet
in
Essette,
Bull.
Trimestriel
Soc.
Mycol.
France
80
('1964'
1965)
Atlas,
pi
147;
Leccinum
lepidum
(Bouchet)
Quadraccia,
Acad.
Naz.
Lincei
264
(1990)
103.
In
previous
studies
(Den
Bakker
et
al.,
2004a, 2004b,
in
press)
we
dealt
with
various
aspects
of
the
evolutionary
ecology
of
the
genus
Leccinum.
The
phylogenetic
results
presented
in
these
studies
also
have
important
taxonomic
implications
on
various
levels,
ranging
from
family
to
species
level.
In
this
chapter
we
will
discuss
these
taxonomic
implications
and
present
a
taxonomic
treatment
of
the
European
representatives
of
the
genus,
with
the
exception
of
two
mediterranean
species,
Leccinum
corsicum
i
1
and
Leccinum
lepidum
2
.
Because
many
species
occur
also
in
North
America
(subsection
Leccinum
-
Den
Bakker
et
al.,
2004b)
or
have
closely
related
sister
species
on
that
continent
(subsection
Scabra
-
Den
Bakker,
submitted),
we
will
also
discuss,
where
possible,
North
American
species
in
additional
notes.

PERSOONIA
-
Vol.
18,
Part
4,
2005512
GENERIC
DELIMITATION
The
original
genus
Leccinum
was
introduced
by
Gray
(1821)
as a
generic
scientific
name
for
boletes
and
contained
species
of
several
currently
recognized
genera,
such
as
Gyroporus,
Boletus,
Suillus,
Chalciporus
and
Xerocomus.
Later
the
use
of
the
name
Leccinum
has
been
limited
to
the
group
of
fungi
we
now
know
as
Leccinum.
Although,
especially
in
temperate
and
boreal
regions,
species
of
the
genus
Leccinum
are
easily
recognized
by
their
prominent,
squamulose
stipe
ornamentation,
the
delimita-
tion
of
the genus
has
been
a
matter
of
discussion
for
decades.
Smith
&
Thiers
(1971)
con-
sidered
the
presence
of
a
squamulose
stipe
ornamentation
that
darkens
with
age
diagnos-
tic
for
the
genus
Leccinum.
Singer (1986)
considered
the
coarse
squamulose
stipe
orna-
mentation
the
most
important
character
of
the
genus
Leccinum,
irrespective
of
colour
or
colour
changes
of
the
squamules.
According
to
Singer (1986)
the
squamules
of
Lec-
cinum
differ
from
those
found
in
other
boletes
by
the
fact
that
the
basidia,
basidioles
and
cystidia
that
make
up
these
squamules
are
positioned
on
a
distinct
hyphal
base,
while
in
other
boletes
they
emerge
directly
from
the
hyphae
in
the
cortex
of
the
stipe.
In
his
discussion
on
the
delimitation
of
the genus
Boletus,
Singer
applied
an
additional
character
to
distinguish
this
genus
from
Leccinum.
Boletes
with
a
yellow
hymenium,
a
squamulose
stipe
and
a
trichodermal
pileipellis
are
considered
to
belong
to
the
genus
Boletus,
while
species
that
share
the
first
two
characters
but
have
either
a
cutis-like
or
epi-
thelial
pileipellis
are
considered
to
belong
to
Leccinum.
Surprisingly,
this last
character
seems
to
be
in
contradiction
with
Singer's
placement
of
J
Leccinum
crocipodium
(L.
nigres-
cens
in
Singer,
1986)
in
Leccinum,
because
a
strict
application
of
his
criteria
would
place
this
species,
which
has
a
trichodermal
pileipellis
and
yellow
hymenium,
in
Boletus.
Sutara
(1989),
in
an
attempt
to
clarify
the
delimitation
of
the genus
Leccinum,
focused
entirely
on
the
anatomy
of
the
stipe
cortex
and
the
stipe
ornamentation.
According
to
Sutara
the
stipe
of
Leccinum
consists
of
longitudinally
positioned
hyphae.
Beneath
the
fertile
layer
composed
of
caulobasidia
and
caulocystidia,
a
thick
(200-1000
pm)
layer
is
present
(the
stipital
lateral
stratum)
over
the
entire
stipe
that
is
composed
of
almost
anticlinally
positioned,
non-interwoven,
parallel
hyphae.
Usually
the
stipital
lateral
stratum
disrupts
at
maturity
of
the
fruit-body,
which
results
in
the
typical
squamulose
stipe
of
Leccinum.
The
Boletus-
type
of
the
stipital
lateral
stratum
is
thin
(20-80(-100)
does
not
rupture
in
mature
fruit-bodies,
its
hyphae
are
not
conspicuously
anticli-
nally
positioned,
its
hyphae
are
often
interwoven
and
in
some
cases
the
stipital
lateral
stratum
is
gelatinized.
Basically
the
anatomical
features
of
the
stipe
lateral
stratum
which
Sutara
described
are
a
more
detailed
description
of
the
anatomical
features
of
the
stipe
that
were
used
by
Singer (1986)
to
distinguish
Leccinum
from
other
genera
in
the
Boletaceae.
Sutara
proposed
putting
all
boletes
with
a
Leccinum-
type
stipital
lateral
stratum
in
Leccinum.
Consequently,
not
only
taxa
classically
referred
to
Leccinum,
but
also
species
like
Boletus
impolitus
3
,
Boletus
depilatus
4
and
Boletus
fragrans
5
belong
in
Leccinum,
if
the
criteria
of
Sutara
are
followed.
3)
Boletus
impolitus
Fr.,
Epicrisis
(1838)
421;
Leccinum
impolitum
(Fr.)
Bertault,
Bull.
Trimestriel
Soc.
Mycol.
France
96
(1980)
287.
4)
Boletus
depilatus
Redeuilh,
Bull.
Trimestriel
Soc.
Mycol.
France
101
('1985'
1986)
396;
Leccinum
depilatum
(Redeuilh)
Sutara,
Ceska
Mykol.
43
(1989)
4.
5)
Boletus
fragrans
Vitt.,
Funghimang.
(1835)
158;
Leccinum
fragrans
(Vitt.)
Sutara,
Ceska
Mykol.
43(1989)
54.

513
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Recently,
molecular
methods
have been used
to
elucidate
relationships
within
the
Boletaceae.
Binder
&
Besl
(2000)
have
used
partial
sequences
of
the
nuclear
ribosomal
large
subunit
(28S
nrDNA)
to
explore
phylogenetic
relationships
between
species
of
Leccinum
and
genera
that
have
traditionally
been
seen
as
transient
to
or
possibly
part
of
Leccinum.
Later
Bresinsky
&
Besl
(2003)
split
off
the
genus
Leccinellum
with
Leccinum
nigrescens
(Richon
&
Roze)
Singer
as
type
species,
based
on
the
results
of
the
study
of
Binder
&
Besl
(2000).
According
to
Bresinsky
&
Besl
this
genus
is
distinguished
from
the
other
species
of
Leccinum
by
the
presence
of
yellow
pigments
in
the
hymenium,
a
blackish
or
greyish
discoloration
of
the
context
of
the
fruit-bodies
when
bruised
and
a
pileus
cuticle
that
consists
of
a
palisade
trichoderm.
Molecular
data
Introduction
and
methods
To
assess
the
actual
support
from
molecular
data
for
the
delimitation
of
Leccinellum
and
the
generic
delimitations
of
Leccinum
as
proposed
by
Smith
and
Thiers
(1971),
Singer (1986)
and
Sutara
(1989),
we
downloaded
a
large
sample
of
28S
sequences
of
Boletales
available
on
GenBank.
We
increased
the
sample
size
to
84 taxa,
as
opposed
to
34
accessions
in
the
original
data
set
of
Binder
&
Besl
(2000).
Not
only
did
we
increase
the
sample
size
of
some
accessions
of
genera
that
have been
considered
closely
related
to
or
even
part
of
Leccinum
(Tylopilus,
Xerocomus,
Boletus
p.p.),
but
we
also
included
the
gasteromycete
genus
Chamonixia,
since
the
results
of
Bruns
et
al.
(1998)
suggested
this
gasteromycete
is
closely
related
to
species
of
Leccinum.
We
are
aware
of
the
fact
that
sequences
submitted
to
Genbank
can
be
subject
to
misidentification
(Bridge
et
al.,
2003;
Vilgalys,
2003)
and
therefore
results
of
a
study
based
on
these
data have
to
be used
with
caution
before
using
these
data
for
taxonomic
changes.
Although
there
is
a
chance
that
part
of
the
tree
is
based
on
wrongly
identified
accessions,
for
which
we
will
provide
likely
examples
below,
we
emphasize
that
re-analyzing
data
that
have
previously
been used
to
propose
taxonomic
changes
can
provide
valuable
insights
in
the
phylogenetic
robustness
of
these
data.
Recently,
Binder
&
Hibbett
(2004)
published
a
tree,
based
on
457
(28S
nrDNA)
sequences
representing
333
species,
of
which
293
belong
to
the
Boletales.
As
several
sequences
are
not
yet
publicly
available,
we
refrain
from
including
this
analysis
in
this
chapter.
We
note,
however,
that
the
conclusions
with
regard
to
Leccinum
as
proposed
here,
do
not
seem
to
need
any
substantial
modification
based
on
this
larger
data
set.
The
downloaded
sequences
were
aligned
using
POA
(Lee
et
al.,
2002;
http://www.
bioinformatics.ucla.edu/poa/POA_Online/Align.html),
and
adjusted
by
eye.
Model-
test
3.4
(Posada
&
Crandall,
1998)
and
MrModeltest
(Nylander,
available
from
http:
//www.ebc.uu.se/systzoo/staff/nylander.html)
were
used
to
determine
the
least
rejected
model
of
sequence
evolution.
The
likelihood
ratio
test
as
implemented
in
Modeltest
and
MrModeltest
(P
>
0.05)
was
used
to
select
the
model
that
was
subsequently
used
in
the
Maximum
Likelihood
and
Bayesian
analyses.
PAUP*4.0bl0
(Swofford,
2002)
was
used
to
perform
the
Maximum
Parsimony
(MP)
and
the
Maximum
Likelihood
analyses,
MrBayes
3.0b4
(Huelsenbeck
&
Ronquist,
2001)
was
used
to
perform
the
Bayesian
analysis.
The
analyses
were
performed
as
described
in
Den
Bakker
(2004b).
MP
bootstrap
support
values
were
calculated
based
on
1000
bootstrap
replicates,
and
1000
trees
were
kept
per
replicate.

514
PERSOONIA
-
Vol.
18,
Part
4,
2005

515
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Results
The
alignment
of
the
sequences
contained
947
sites,
of
which
332
were
variable
and
229
were
parsimony-informative.
The
MP
analysis
resulted
in
1902
MP
trees
of
1335
steps
(C.I.
=
0.367,
R.I.
=
0.687).
Based
on
the
outcome
of
Modeltest
the
Tamura-Nei
(Tamura
&
Nei,
1993)
model
with
invariable
sites
and
variable
sites
following
a
gamma
distribution
was
chosen.
For
this
analysis
the
following
settings
were
used:
base
fre-
quencies
A:
0.2789,
C:
0.2089,
G:
0.2769,
T:
0.2344;
substitution
rates
A«C:
1,
A«G:
2.9695,
A-m-T:
1,C**G:
1,C«*T:
8.1475;
G*-*T:
1;
assumed
proportion
of
invariable
sites
=
0.4761;
shape
parameter
a
=
0.4828.
The
ML
analysis
resulted
in
one
tree
of
-In
L
score
8021.60869.
For
the
Bayesian
analysis
the
general-time
reversible
model
(Rodriguez
et
al.,
1990)
was
used.
Fig.
1
shows
a
phylogram
in
which
the
results
of
the
phylogenetic
analyses
are
sum-
marized.
With
respect
to
the
position
of
Leccinum
the
trees
obtained
from
the
different
analyses
did
not
differ.
Almost
all
species
that
were
placed
in
Leccinum
by
Singer
(1986)
are
found
in
a
well-supported
clade,
except
for
Boletus
eximius
6
.
Boletus
eximius,
a
species
that
has
a
strongly
squamulose
stipe
ornamentation,
differs
from
species
of
Leccinum
by
the
overall
dark
purplish
colour
of
its
fruit-body.
Remarkably,
the
branches
that
form
the
Leccinum
clade
are
all
long
in
comparison
to
other
Boletales
in
the
phylogram,
which
could
indicate
that
some
of
the
accessions
are
wrongly
placed
in
Leccinum
because
of
a
phenomenon
called
long-branch
attraction
(Felsenstein,
2004).
Mainly
phylogenies
based
on
a
MP
optimalisation
are
susceptible
to
this
phenomenon
(Felsenstein,
2004).
However,
the
fact that
more
or
less
the
same
topology
is
also
recovered
under
ML
and
Bayesian
optimalisation
criteria
pleas
against
long-branch
attraction.
Although
the
genus
Leccinum
is
represented
as a
well-supported
clade,
28S
seems
to
lack
sufficient
information
to
resolve
the
phylogeny
of
the
Boletales.
A
putative
sistergroup
to
Leccinum
can
therefore
not
be
assigned.
According
to
this
phylogram
Leccinum
is
a
north-temperate
genus
with
several
species
in
Australasia
(mainly
in
sect.
Roseoscabra
i)
and
Central
America.
Various
species
in
Leccinum
have been
described
from
tropical
Africa
(Heinemann,
1964),
but the
delimitation
from
tropical
African
species
of
Tylopilus
in
still
unclear
(Th.W.
Kuyper,
pers.
comm.).
No
molecular
data
are
available
of
these
species
and
their
position
can
therefore
not
be
assessed.
A
phylo-
genetic
analysis
of
tropical
African
species
that
have been
described
in
Leccinum
and
Tylopilus
is
urgently
needed.
Discussion
Morphological
distinction
of
Leccinum
from
other
boletes
seems
to
be
difficult,
even
if
we
omit
the
recently
derived
sequestrate
genera
Chamonixia
and
Octavianina.
6)
Boletus
eximius
Peck,
J.
Mycol.
3
(1887)
54;
Tylopilus
eximius
(Peck)
Singer,
Am.
Midi.
Naturalist
37
(1947)
109;
Leccinum
eximium
(Peck)
Singer,
Persoonia
7
(1973)
319.
Fig.
1
(Opposite
page).
28S
nrDNA
phylogram
with
the
highest
posterior
probability
as
inferred
by
Bayesian
analysis.
Thickened
branches
receive
posterior
probabilities
of
≥
95%.
Branches
marked
with
a
asterisk
receive
a
MP
bootstrap
support
≥
70%.
Taxa
marked
with
an
asterisk
have
been
proposed
to
belong
to
the
genus
in
the
past.
Leccinum

PERSOONIA
-
Vol.
18,
Part
4,
2005516
In
Europe
species
of
the
genus
Leccinum
can
be
recognized
by
the
combination
of
the
following
characters:
a
squamulose
stipe
ornamentation,
pores
that
are
either
brownish,
whitish
or
yellowish
(not
reddish
as
in
Boletus
erythropus
7
),
and,
if
the
hymenium
contains
yellowish
pigments,
the
context
of
the
stipe
usually
discolours
greyish
or
blackish
when
bruised.
According
to
the
phylogram
depicted
in
Fig.
1,
L.
chromapes
8
is
sister
to
the
remainder
of
Leccinum
plus
Chamonixia.
Its
taxonomic
position
is
still
doubtful.
Hailing
&
Mueller
(2003)
considered
it
a
member
of
Tylopilus,
at
the
same
time
treating
another
species
closely
resembling
it
as
Leccinum
cartagoense
9
.
Curiously,
in
the
cladogram
produced
by
Binder
&
Hibbett
(2004)
Tylopilus
chromapes
(AF139709)
belongs
to
the
Leccinum
clade,
while
accession
AY612834
is
placed
in
a
clade
with
various
other
Tylopilus
species.
Evidently,
the
taxonomic
identity
of
both
accessions
needs
to
be
determined
before
the
position
of
L.
chromapes
can
be
elucidated.
The
sequestrate
(truffle-like)
Chamonixia
caespitosa
7
10
is
nested
within
Leccinum.
In
the
MP
analysis
it
is
found
in
a
strongly
supported
clade
with
amongst
others
L.
corsicum,
L.
crocipodium
and
i
L.
pseudoscabrum.
The
ML
and
Bayesian
analyses
do
not
support
this
placement
and
place
Chamonixia
basal
to
the
clade
containing
sections
Leccinum
and
Luteoscabra.
Analyses
of
mtDNA
(Bruns
et
al.,
1998)
and
chemotaxonomic
evidence
(the
presence
of
the
cyclopentenones
gyrocyanin
and
gyroporin)
is
consistent
with
such
a
placement
(Gill
&
Steglich,
1987).
Remarkably
this
is
not
the
only
sequestrate
genus
that
is
derived
from
a
Leccinum-like
ancestor.
Binder
(1999)
and
Binder
&
Hibbet
(2004)
showed
(based
on
the
phylogenetic
analysis
of
28S-sequences)
that
Octavianina
is
also
nested
within
Leccinum.
These
sequences
are
at
present
not
yet
available
on
Genbank,
making
it
impossible
to
reassess
their
phylogenetic
position.
The
phylogenetic
position
of
these
sequestrate
genera
implies
that
the
evolutionary
step
from
a
normal
fruit-body
to
a
truffle-like
fruit-body
is
a
relatively
small
one.
Evidence
for
such
rapid
evolutionary
transitions
from
boletoid
and
agaricoid
ancestors
to
sequestrate
taxa
is
not
uncommon
(Bruns
et
al.,
1989;
Peintner
et
al.,
2001;
Miller
et
al.,
2001;
Binder
&
Bresinsky,
2002).
Interestingly,
both
Chamonixia
and
Octavianina
have
ornamented
spores,
while
Leccinum
has
without
exception
smooth
spores.
A
taxonomical
consequence
of
the
phylogenetic
placement
of
Chamonixia
(and
maybe
Octavianina)
in
the
Leccinum
clade
could
be
to
sink
these
sequestrate
taxa
in
Leccinum.
Other
options
would
be
to
remove
L.
chromapes
from
Leccinum
or
to
subdivide
the
genus
Leccinum
into
smaller
genera.
However,
lack
of
sufficient
resolution
of
the
cladogram
and
different
results
from
the
MP,
ML
and
Bayesian
analyses
make
it
hardly
recommendable
to
propose
taxonomic
changes
for
the
time
being.
Apart
from
Chamonixia,
Leccinum
consists
of
two
clades
that
have been
known
as
sect.
Luteoscabra
and
sect.
Leccinum.
The
MP
analysis
showed
high
bootstrap
support
7)
Boletus
erythropus
Pers.,
Observ.
mycol.
1
(1796)
23
:
Fr.;
Boletus
luridus
8
[var.]
erythropus
(Pers.:
Fr.)
Fr.,
Syst.
mycol.
1
(1821)
391.
8)
Boletus
chromapes
Frost,
Bull.
Buffalo
Soc.
Nat.
Sci.
2
(1874)
105;
Tylopilus
chromapes
(Frost)
A.H.
Sm.
&
Thiers,
Boletes
Michigan
(1971)
92;
Leccinum
chromapes
(Frost)
Singer,
Am.
Midi.
Naturalist.
37
(1947)
124.
9)
Tylopilus
cartagoensis
Wolfe
&
Bougher,
Austral.
Syst.
Bot.
6
(1993)
191;
Leccinum
cartagoense
(Wolfe
&
Bougher)
Hailing
&
G.M
Muell.
in
Hailing,
Kew Bull.
54
(1999)
747.
10)
Chamonixia
caespitosa
Rolland,
Bull.
Soc.
Mycol.
France
15
(1899)
73-78.

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
517
(77%)
for
the
clade
that
largely
coincides
(if
we
do
not
consider
the
sequestrate
genera)
with
section
Luteoscabra
Singer
(1947).
The
posterior
probability
of
this
clade
is
less
than 90
%,
which
may
indicate
that
the
strong
MP
bootstrap
support
might
be
caused
by
long-branch
attraction
and
may
falsely
suggest
a
monophyletic
(natural)
group.
The
only
difference
with
the
original
section
Luteoscabra
is
that,
as
already
indicated
by
Lannoy
&
Estades
(1995),
species
like
L.
pseudoscabrum
and
L.
albellum
11
also
belong
to
this
clade.
Bresinsky
&
Besl's
genus
Leccinellum
overlaps
with
the
Luteoscabra
clade
found
in
this
analysis,
consisting
of
L.
pseudoscabrum,
L.
albellum
and
L.
cro-
cipodium
and
related
species,
but
for
unclear
reasons
they
exclude
L.
subglabripes
12
,
L.
rubropunctum
13
and
‘Boletus’
longicurvipes
14
from
Leccinellum,
thereby
leaving
the
genus
Leccinellum
paraphyletic.
Also
from
a
morphological
point
of
view
we
do
not
think
there
are
grounds
for
assuming
that
Leccinellum
as
defined
by
Bresinsky
&
Besl
forms
a
monophyletic
group
that
can
be
discriminated
from
other
boletes
by
one
or
more
diagnostic
characters.
Moreover,
the
28S
data
seem
to
be
insufficient
or
even
contradictory
for
the
recognition
of
Leccinellum
as a
distinct
genus.
Taking
monophyly
of
a
genus
as
an
essential
criterion
for
generic
recognition,
our
delimitation
of
the
genus
will
be
more
or
less
congruent
with
that
of
Singer
(1986).
The
group
of
North
American
species
(B.
longicurvipes,
B.
rubropunctus,
B.
sub-
glabripes
and
B.
hortonii
15
in
the
analysis)
that
has
previously
been
classified
as
Boletus
section
Pseudoleccinum
by
Smith
&
Thiers
(1971)
seems
to
pose
a
problem.
Species
of
this
section
have
a
squamulose
stipe
ornamentation,
a
yellowish
hymenium,
but
do
not
show
the
blackish/greyish
discoloration
of
the
context
that
is
found
in
European
species
of
Leccinum
with
a
yellowish
hymenium.
The
28S
analysis
indicates
that
B.
longicurvipes,
B.
rubropunctus,
and
B.
subglabripes
are
well
nested
within
Leccinum
and
should
therefore
be
considered
species
of
Leccinum.
Boletus
hortonii
on
the
other
hand,
even
though
morphologically
very
close,
is
found
in
a
clade
together
with
spe-
cies
of
Xerocomus,
B.
impolitus
and
B.
depilatus.
While
molecular
data
put
most
of
the
species
of
section
Pseudoleccinum
in
Leccinum,
chemotaxonomic
data
(the
presence
of
the
pulvinic
acid
derivatives
xerocomic
acid
and
variegatic
acid)
suggest
a
relation-
ship
with
Boletus
and
not
with
Leccinum
(Gill
&
Steglich,
1987).
To
what
extent
these
molecular
and
chemotaxonomic
results
are
influenced
by
misidentifications
and
other
errors
in
sequence
databases
remains
to
be
investigated.
INFRAGENERIC
DELIMITATION
Given
the
generic
delimitation
discussed
above,
the
three
main
monophyletic
groups
correspond
largely
to
the
subdivision
in
sections
by
Singer
(1986),
being
(1)
section
11)
Boletus
albellus
Peck,
Rept.
N.Y.
State
Mus.
41
(1889)
149;
Leccinum
albellum
(Peck)
Singer,
Mycologia
37(1945)799.
12)
Boletus
subglabripes
Peck,
Bull.,
New
York
State
Mus.
Nat.
Hist.
8
(1889)
112;
Leccinum
subglabripes
(Peck)
Singer,
Mycologia
37
(1945)
799.
13)
Boletus
rubropunctus
Peck,
Rep.
New
York
State
Mus.
Nat.
Hist.
50
(1898)
109;
Leccinum
rubropunctus
(Peck)
Singer,
Amer.
Midi.
Naturalist.
37
(1947)
117.
14);
Boletus
longicurvipes
Snell
&
A.H.
Sm.,
J.
Elisha
Mitchell
Sci.
Soc.
56
(1940)
325.
15)
Boletus
hortonii
A.H.
Sm.
&
Thiers,
Boletes
Michigan
(1971)
319;
Leccinum
hortonii
(A.H.
Sm.
&
Thiers)
Hongo
&
Nagas.,
Rep.
Tottori
Mycol.
Inst.
16
(1978)
50.

PERSOONIA
-
Vol.
18,
Part
4,
2005518
Roseoscabra
(type
species
L.
chromapes
;)
-
if
indeed
best
classified
in
Leccinum,
(2)
section
Luteoscabra
(type
species
L.
nigrescens
=
L.
crocipodium
i)
and
(3)
section
Lec-
cinum
(type
species
L.
aurantiacum
i).
In
Europe
especially
section
Luteoscabra
and
Leccinum
are
important,
species
of
section
Roseoscabra
are
mainly
found
in
Australasia
and
North
and
Central
America
(Wolfe
&
Bougher,
1993;
Hailing
&
Mueller,
2003).
In
the
infrageneric
classification
of
Smith
et
al.
(1967)
and
Lannoy
&
Estades
(1995),
section
Roseoscabra
is
lacking,
either
because
it
is
considered
a
section
of
Tylopilus,
as
is
the
case
in
Smith
et
al.
(1967),
or
simply
because
the
subdivision
is
completely
based
on
the
European
taxa
as
in
Lannoy
&
Estades
(1995).
Taking
into
consideration
the
molecular
phylogenetic
results
of
Den
Bakker
et
al.
(2004a,
b),
three
subclades
can
be
recognized
within
section
Leccinum
and
these
are
treated
here
as
subsections.
This
infrageneric
subdivision
will
also
be
followed
in
the
treatment
of
the
genus
in
the
Flora
agaricina
neerlandica:
(1)
Subsection
Leccinum.
Pileus
margin,
especially
in
young
fruit-bodies,
consisting
of
irregularly
disrupted
flaps
(see
Fig.
5).
Usually
species
in
this
group
show
a
blackish
or
greyish
discoloration
of
the
context
when
bruised.
(2)
Subsection
Fumosa
A.H.
Sm.,
Thiers
&
Watling.
Pileus
margin
entire
(see
Fig.
10),
context
usually
discolouring
greyish
when
bruised.
Note,
however,
that
according
to
our
observations
this
reaction
can
be
present
or
absent
in
individual
fruit-bodies
of
the
same
species.
Generally
species
of
this
subclade
are
associated
with
species
of
Populus.
(3)
Subsection
Scabra
Pilat
&
Dermek.
Pileus
margin
entire,
greyish
or
blackish
dis-
coloration
of
the
context
is
lacking.
According
to
our
data
species
of
this
section
are
exclusively
associated
with
Betula,
though
some
authors
(Lannoy
&
Estades,
1995)
claim
that
some
species
can
also
be
associated
with
Salix.
SPECIFIC
DELIMITATION
In
the
Flora
agaricina
neerlandica
a
strictly
morphological
species
concept
is
used
(Kuyper,
1988).
In
this
sense
Van
Steenis'
(1957)
statement
is
followed:
a
'good'
species
differs
in
at
least
two,
independent
morphological
characters.
However,
from
Kuyper's
essay
it
can
be
deduced
that
we
are
not
dealing
with
a
typological
view,
but
that
actually
the
use
of
the
morphological
species
concept
should
be
seen
as a
hypothesis
of
what
the
boundaries
of
'natural'
species
(biological
(Mayr,
1957)
or
evolutionary
species
(Simpson,
1951;
Wiley,
1978)
are.
Here
we
use
the
term
'natural'
species
for
species
as
individuals
in
a
philosophical
sense,
i.e.
we
assume
that
species
exist
in
nature,
independent
of
our
ability
to
recognize
them,
and
that
species
are
able
to
evolve
and
speciate.
This
is
in
contrast
to
a
strict
typological
species
concept
where
species
are
more
or
less
considered
natural
kinds,
constructions
of
the
human
mind,
without
any
necessary
real
existence,
and
evolution
of
the
taxon
is
only
possible
when
this
results
in
a
new
class
(species).
Because
we
are
not
only
interested
in
recognizing
species,
but
also
in
their
ecology
and
their
evolutionary
history,
we
consider
species
individuals.
As
such
we
see a
taxonomy
based
on
morphology
as a
hypothesis
of
what
the
boundaries

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
519
morphological
criteria
(Lannoy
&
Estades,
1995,
Smith
&
Thiers,
1971)
to
the
left, and
to
the
right
the
accepted
names
are
given
as
a
result
of
the
current
species
concept.
Thickened
branches
receive
posterior
probabilities
of
≥ 95%.
Grey
thickened
branches
receive
posterior
probabilities
between
90-95%.
Values
above
clades
indicate
MP
bootstrap
values,
values
on
the
left
side
of
the
slash
indicate
the
bootstrap
value
calculated
when
partial
sequences
are
included,
on
the
right
site
bootstrap
values
when
the
partial
sequences
are
excluded
(see
Den
Bakker
et
al.,
2005).
Bootstrap
values
<
60%
are
not
indicated.
Type-accessions
are
printed
bold.
data.
Individual
accessions
are
named
with
the
traditional
Gapdh
Fig.
2.
One
of
84
ML
trees
based
on

PERSOONIA
-
Vol.
18,
Part
4,
2005520
of
species
as
individuals
are.
Additional
information
from
for
instance
breeding
experi-
ments
and
molecular
phylogenies
can
be used
to
adjust
the
hypothesis
about
boundaries
of
species.
Combining
biological
(breeding
experiments),
evolutionary
(molecular
phylogenies)
and
phenetic
(morphology)
data
in
one
species
concept
is
by
no means
easy.
Aanen
&
Kuyper
(2004)
described
their
approach
in
arriving
at
an
operational
species
concept
in
the
Hebeloma
crustuliniforme
complex.
Their
approach
involved
the
use
of
a
phenetic
Fig.
3.
ML
tree
based
on
ITS2
sequences.
Individual
accessions
are
named
with
the
traditional
morphological
criteria
(Lannoy
&
Estades,
1995,
Smith
&
Thiers,
1971)
to
the
left,and
at
the
right
the
accepted
names
are
given
as
a
result
of
the
current
species
concept.
Thickened
branches
receive
posterior
probabilities
of
≥
95%.
Grey
thickened
branches
receive
posterior
probabilities
between
90
and
95%.
Values
above
clades
indicate
MP
bootstrap
values,
values
<
50%
are
not
indicated.
Type-
accessions
are
printed
bold.

521
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
concept
that
is
at
least
consistent
with
biological
criteria
(intercompatible
collections
should
not
be
classified
as
different
species)
and
evolutionary
criteria
(species
that
turn
out
to
be
polyphyletic
should
be
rejected).
In
the
case
of
Leccinum,
a
similar
approach
to
arrive
at
consistency
between
phenetic
and
evolutionary
data
was
used.
It
is
important
to
reassess
the
status
of
morphospecies
if
molecular
data
show
that
the
morphospecies
is
clearly
polyphyletic.
Basically
two
classes
of
explanations
for
polyphyletic
morphospecies
can
be distin-
guished.
The
first
class
of
explanations
are
biological
causes
for
polyphyly,
like
hybridi-
sation.
The
second
class
of
explanations
is
that
we
are
actually
dealing
with artificial
species
(natural
kinds)
and that
the
morphological
character
states
that
are
considered
diagnostic
for
such
a
taxon
fall
within
the
phenotypic
variability
of
more
than
one
spe-
cies.
Careful
reassessment
of
other
morphological
characters
usually
shows
that
other
characters
are
indicative
for
the
monophyletic
groups
that
can
be
considered
species.
Regarding
the
names
of
these
groups,
type
specimens
were
used
as
reference
points
and
if
more
than
one
type
specimen
was
present
in
a
monophyletic
species,
the
classical
nomenclatural
rules
with
respect
to
priority
and
synonymy
of
names
were
applied.
Practical
application
The
practical application
of
the
operational
species
concept
as
practised
here
will
be
illustrated
with
the
examples
of
L.
scabrum,
L.
holopus
and
L.
pulchrum
and
two
gene
trees
(Figs.
2,
3).
In
these
trees
the
individual
accessions
are
named
on
the
basis
of
traditional
morphological
criteria
(Lannoy
&
Estades,
1995)
to
the
left,
and
to
the
right
the
accepted
names
are
given
as a
result
of
the
current
species
concept.
Leccinum
scabrum
Traditionally,
L.
scabrum
is
considered
a,
Betula-
associate
with
a
blackish
stipe
orna-
mentation
and
a
brownish
pileus.
The
most
important
diagnostic
criterion,
however,
is
the
fact
that
the
context
does
not
change
colour
when
bruised
or
becomes
at
most
a
little
pinkish.
The
gene
trees
(Figs.
2,3)
show
that
accessions
named
L.
scabrum
are
found
in
three
well-supported
clades
in
the
Gapdh
tree
(the
L.
snellii
'
16
clade,
the
L.
schis-
tophilum
and
the
L.
scabrum/L.
melaneum
clade)
and
in
one
well-supported
clade
(L.
snellii
clade)
and
one
weakly
supported
clade
(L.
scabrum/L.
rotundifoliae)
in
the
ITS2
tree.
As
indicated
above,
the
polyphyly
of
L.
scabrum
can
mean
two
things,
either
L.
sca-
brum
is
an
artificial
species
or
L.
scabrum
is
a
'real'
taxonomic
entity
and
the
characters
that
are
generally
considered
diagnostic
for
the
species
need
to
be
reconsidered.
Of
course
there
is
a
trivial
third
alternative
explanation,
viz.
that
the
whole
clade
constitutes
just
one
very
variable
species.
However,
patterns
of
morphological
differences,
cor-
related
with
molecular
divergence,
militate
against
that
solution.
In
the
case
of
L.
sca-
brum
the
second
option
was
chosen.
The
accessions
that
were
named
L.
scabrum
but
were
placed
in
the
L.
snellii
and
L.
schistophilum
clade
were
not
microscopically
differ-
ent
from
accessions
of
these
species
that
showed
a
typical
(bluish
and
pinkish)
discolora-
tion
of
the
context.
Therefore
it
can
be
concluded
that
characters
based
on
discoloration
of
the
context
cannot
be used
to
discriminate
L.
scabrum
from
some
(atypical)
forms
16
Leccinum
snellii
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
6
(1967)
120,
Figs.
3,4.

PERSOONIA
-
Vol.
18,
Part
4,
2005522
of
other
species.
Morphological
comparison
of
accessions
of
the
L.
snellii,
L.
schis-
tophilum
and
the
L.
scabrum/L.
melaneum
clade
in
the
Gapdh
tree
showed
that
mainly
microscopical
characters
(pileipellis
structure,
spore
shape
and
caulocystidia)
can
be
used
to
distinguish
accessions
of
these
clades
from
each
other
(see
key).
Leccinum
holopus
Both
the
Gapdh
(Fig.
2)
and
the
ITS2
gene
tree
(Fig.
3)
show
a
monophyletic
clade
for
L.
holopus
and
both
trees
suggest
that
the
eastern
North
American
accessions
form
a
monophyletic
group
on
their
own.
Morphologically,
however,
there
is
no
character
that
can
be used
to
tell
accessions
of
the
North
American
clade
apart
from
European
collections,
and
therefore,
because
basically
a
morphological
species
concept
is
used,
this
clade
cannot
be
considered
a
species
on
its
own.
Leccinum
pulchrum
Specimens
identified
as
L.
pulchrum
are
found
in
three
monophyletic
clades
in
the
Gapdh
tree:
one
accession
is
in
the
L.
scabrum
clade,
the
accessions
of
the
type
material
are
found
in
the
L.
rotundifoliae
clade
and
one
accession
designated
as
L.
pulchrum
is
found
in
the
L.
holopus
clade.
For
this
reason
we
see
L.
pulchrum
as
circumscribed
by
Lannoy
&
Estades
(1995)
as
an
artificial
taxon.
The
morphological
characters
(mainly
the
pileus
colour
and
the
discoloration
of
the
context
when
bruised)
on
which
the
identification
of
L.
pulchrum
is
based
are
apparently
not
diagnostic
for
a
molecular
monophyletic
taxon.
Based
on
the
sequence
data
L.
pulchrum
has
to
be
synonymized
with
L.
rotundifoliae,
a
boreal,
subalpine
and
arctic
taxon
that
has
been
confused
with
L.
holopus
in
the
past
(hence
the
presence
of
some
L.
rotundifoliae
accessions
in
the
L.
holopus
clade
in
the
Gapdh
and
ITS2
gene
trees).
These
results
show
that
the
morpho-
logical
variability
of
L.
rotundifoliae
is
wider
than
previously
understood.
Re-examina-
tion
of
the
L.
pulchrum
accessions,
on
the
basis
of
a
set
of
different
characters,
in
the
L.
holopus
and
the
L.
scabrum
clade
showed
that
these
fit
the
morphological
concept
of
these
species,
and
must
be
considered
as
misidentifications.
INFRASPECIFIC
DELIMITATION
Most
species
that
are
delimited
according
to
the
species
concept
that
is
previously
discussed
show
a
continuous
variability
of
morphological
characters
like
pileus
colour
and
discoloration
of
the
context.
In
L.
holopus
we
observe
a
different
pattern.
In
this
taxon
fruit-bodies
that
are
found
in
Sphagnum
bogs
or
other
wet
acidic
environments
have
the
typical
slender
habitus
of
L.
holopus,
have
pale
whitish
stipe
squamules
and
do
not
show
any
noticeable
discoloration,
except
for
some
tiny
bluish
spots
in
the
stipe
base
or
the
basal
mycelium.
Fruit-bodies
of
the
same
species
found
in
drier
habitats,
like
marshy
forest,
usually
have
a
less
slender
habitus
(comparable
to
L.
scabrum
0,
brownish
to
blackish
stipe
squamules
and
pinkish
to
reddish
discoloration
of
the
stipe
context
and
a
green
bluish
discoloration
in
the
stipe
base.
In
Europe
the
latter
form
has
been
described
as
L.
nucatum
Lannoy
&
Estades.
Microscopically
the
two
forms
are
not
distinct,
neither
are
they
in
the
Gapdh
and
ITS2
trees.
Moreover
in
Europe
inter-
mediates
between
L.
holopus
and
L.
nucatum
can
be
found,
and
therefore
we
do
not
consider
L.
nucatum
of
any
taxonomic
value.
The
situation
in
North
America
seems
to
be
different.
Here
we
find
apart
from
the
normal
slender,
whitish
form
of
L.
holopus,

523
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
L.
holopus
var.
americanum
1
17
Smith
&
Thiers
(see
discussion
under
L.
holopus
for
diag-
nostic
characters
of
L.
holopus
var.
americanum
).
Both
gene
trees
indicate
that
there
is
a
geographical
distinction
between
accessions
of
L.
holopus
from
eastern
North
America
and
from
Europe
and
the
arctic
regions.
However,
there
is
no
relation
with
the
varieties
recognized.
In
North
America,
the
varieties
seem
to
be
rather
constant
throughout
the
distribution
area.
We
therefore
think
it
is
legitimate
to
recognize
them
taxonomically.
HYBRIDISATION
AND
INTROGRESSION
Two
taxa
seem
to
pose
a
serious
problem
when
we
apply
the
criterion
of
monophyly
in
at
least
one
gene
tree
without
this
monophyly
being
significantly
contradicted
by
other
gene
trees.
The
first
problematic
taxon
is
L.
cyaneobasileucum
(as
L.
brunneo-
griseolum
in
Den
Bakker,
2005).
This
taxon
is
morphologically
distinct
from
L.
holopus
in
most
of
its
microscopical
and
macroscopical
characters.
The
Gapdh
tree
shows
that
accessions
of
this
species
form
a
highly
supported
monophyletic
clade
and
indicates
that
this
species
diverged
from
L.
holopus
long
before
a
divergence
of
European
and
North
American
L.
holopus
took
place.
Moreover,
the
divergence
between
L.
holopus
and
L.
cyaneobasileucum
(1.7
to
2.3
%
divergence)
is
comparable
to
the
divergence
between
L.
scabrum
and
L.
rotundifoliae
(1.5
to
2.3
%
divergence),
which
indicates
that
L.
holopus
and
L.
cyaneobasileucum
diverged
a
long
time
ago,
given
the
factthat
Gapdh
evolves
clock-like
in
Leccinum
subsect.
Scabra
(Den
Bakker,
2005).
The
ITS2
tree,
however,
shows
that
L.
cyaneobasileucum
is
placed
in
the
European/arctic
clade
of
L.
holopus.
Other
studies
(Binder,
1999;
Den
Bakker
et
al„
2004a)
show
that
the
sequence
of
ITS
1
of
this
species
is
also
identical
to
that
of
European
accessions
of
L.
holopus.
Considering
the
morphological
distinctness
of
L.
cyaneobasileucum
and
the
fact that
Gapdh
seems
to
indicate
that
this
taxon
diverged
from
L.
holopus
a
long
time
ago,
we
consider
this
taxon
a
species
withminimal
phylogenetic
quality.
Probably
recent
hybridization
between
L.
cyaneobasileucum
and
L.
holopus
and
subsequent
introgres-
sion
has
resulted
in
the
introduction
of
an
L.
holopus-
-ITS
in
the
L.
cyaneobasileucum
genome.
A
second
problem
is
L.
melaneum.
Collections
of
this
species
differ
mainly
from
L.
scabrum
in
the
greyish
colour
of
the
stipe.
In
other
macroscopical
and
microscopi-
cal
characters
it
is
identical
to
L.
scabrum.
The
ITS2
tree
shows
that
L.
melaneum
accessions
form
a
well-supported
clade
of
their
own.
Visual
comparison
of
the
ITS2
sequences
with
other
sequences
of
species
of
subsection
Scabra
showed
that,
besides
several
autapomorphic
character
states,
a
deletion
that
is
present
in
L.
scabrum
and
L.
rotundifoliae,
but
absent
in
other
species
of
subsection
Scabra
is
also
absent
in
L.
melaneum.
Given
the
fact that
L.
melaneum
shows
several
molecular
autapomorphic
characters,
we
think
that
L.
melaneum
represents
an
ancient
hybrid
between
L.
scabrum
and
a
L.
holopus-like
ancestor.
The
molecular
data
indicate
that
this
taxon
is
evolving
independently
from
L.
scabrum.
Strictly,
if
we
were
to
apply
the
two
character
rule
to
L.
melaneum,
we
would
have
to
consider
it
a
variety
of
L.
scabrum.
However,
given
its
putative
hybridogenic
origin
we
prefer
to
give
it
specific
status,
until
further
molecular
research
has
resolved
its
taxonomic
status.
11)
Leccinum
holopus
var.
americanum
A.H.Sm.
&
Thiers,
Boletes
Michigan
(1971) 183

PERSOONIA
-
Vol.
18,
Part
4,
2005524
REVISION
OF
LECCINUM
MATERIAL
AND
METHODS
Macroscopical
and
ecological
characters
The
discussion
of
the
macroscopical
and
ecological
characters
is
mainly
based
on
personal
observations
of
the
authors;
the
descriptions
of
the
individual
species
are
based
on
our
own
observations,
sometimes
supplemented
with
descriptions
from
literature
or
fieldnotes
and
descriptions
of
other
mycologists.
Microscopical
characters
The
pileipellis
was
studied
in
radial
section
of
the
pileipellis
and
mounted
in
water.
All
observations
that
are
discussed
are
made
in
this
way
unless
indicated
otherwise.
Spores
were
mounted
in
demineralised
water
and
observed
under
oil-immersion.
Only
spores
from
the
(pileal)
hymenium
were
used
and
an
attempt
was
made
to
measure
only
mature
spores.
Spores
were
considered
mature
when
a
clearly
developed,
slightly
(brownish)
coloured
spore
wall
was
present
and
guttules
could
be
observed
within
the
spore.
Sometimes
extremely
elongate
spores
(Q
>
4)
were
present
in
the
hymenium
of
older
fruit-bodies.
These
are
considered
anomalies,
probably
associated
with
unnatural
aging
of
the
fruit-bodies.
These
were
not
included
in
the
measurements.
Circa
30
spores
per
collection
were
measured.
Hymenocystidia
were
observed
in
a
solution
of
5%
KOH.
To
observe
caulocystidia
a
stipital
squamule
was
picked
from
halfway
the
stipe
with
a
pair
of
fine
tweezers,
mounted
in
a
5%
KOH
solution
and
squashed.
MACROSCOPICAL
CHARACTERS
Discoloration
of
context
In
many
species
the
context
of
the
pileus
and
stipe
changes
colour
when
exposed
to
theair.
Traditionally
the
discoloration
of
the
context
has
been
considered
an
important
character
in
the
classification
of
Leccinum.
The
most
important
discolorations
of
the
usually
white
context
are
a
blackish,
a
bluish
and/or
a
pinkish
to
reddish
discoloration.
Especially
the
pinkish-reddish
and
blackish
discolorations
of
the
context
are
often
eas-
ily
observed
when
a
fruit-body
is
cut
in
half
and
the
exposed
surface
is
bruised
with
a
blunt
object,
for
instance
the
blunt
side
of
a
knife.
Usually
the
discoloration
starts
within
seconds
after
the
context
is
bruised,
however,
the
bluish
discoloration
can
sometimes
appear
after
several
hours
and
is
sometimes
only
visible
in
parts
of
the
fruit-body
that
have
already
been
damaged
by
snails
or
arthropods.
The
absence
of
a
blackish
discoloration
has
been
used
by
Lannoy
&
Estades
(1995)
as a
diagnostic
character
of
subsection
Scabra.
Although
our
ITS
results
showed
that
subsection
Scabra,
if
defined
by
this
character,
is
probably
polyphyletic,
Gapdh-
data
show
considerable
support
for
the
monophyly
of
subsection
Scabra
as
defined
by
Lan-
noy
&
Estades
(1995).
At
the
species
level
the
intensity
of
the
blackish
discoloration
seems
to
help
with
species
identification
within
subsection
Leccinum.
The
context
of
L.
vulpinum
darkens
generally
less
intensively
than
that
of
related
species
such
as
L.
insigne
18
,
L.
aurantiacum
and
L.
versipelle.
18),
Leccinum
insigne
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
5
(1966)
160.
Fig.
13.

525
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
A
pinkish,
sometimes
reddish,
discoloration
of
the
context
is
considered
to
be
a
diagnostic
character
for
species
within
subsections
Scabra
and
Leccinum.
Smith
&
Thiers
(1971)
made
a
distinction
in
their
keys
between
species
in
subsection
Leccinum
that
show
a
pinkish
discoloration
before
turning
blackish
(like
L.
aurantiacum
sensu
Smith
&
Thiers)
and
those
that
do
not
show
a
pinkish
discoloration,
but
only
a
blackish
discoloration
(like
L.
insigne
>).
We
found
that
this
pinkish
discoloration
was
often
lacking
in
species
that
were
supposed
to
have
it
according
to
literature
and
found
that
usually
the
European
species
can
be
recognized
by
other,
more
stable
characters.
Watling
(1970)
and
Lannoy
&
Estades
(1995)
considered
the
intensity
of
the
pinkish
discoloration
a
distinctive
character
to
distinguish
L.
scabrum
and
related
species.
Leccinum
scabrum
is
supposed
to
display
no
or
only
a
slightly
pinkish
discoloration,
while
species
like
L.
roseofractum
are
supposed
to
show
a
rapid
reddish
discoloration.
We
found
samples
identified
as
L.
scabrum
and
L.
roseofractum
in
one
clade
and
could
not
find
any
phy-
logenetic
signal
that
these
were
two
distinct
species.
Moreover,
we
found
that
the
inten-
sity
of
the
pinkish
discoloration
of
the
context
of
the
accessions
in
this
clade
is
a
gradual
one
and
can
therefore
not
be used
as a
diagnostic
character
of
any
of
these
spe-
cies.
A
bluish
discoloration
is
present
in
most
species
of
subsections
Leccinum,
Fumosa
and
Scabra,
except
in
L.
scabrum
and
L.
rotundifoliae.
This
blue
discoloration
is
usually
found
in
the
cortex
of
the
stipe
base
and/or
in
the
basal
mycelium,
though
in
some
species
like
L.
variicolor,
it
is
found
in
the
cortex
of
the
lower
half
of
the
stipe.
The
colour
is
usually
blue
(K.
&
W.
23A7)
or
greenish
blue
(24A5,25
A5).
In
L.
variicolor
the
greenish
blue
discoloration
usually
changes
to
yellow
in
dried
fruit-bodies,
a
phenomenon
rarely
observed
in
other
species
of
subsection
Scabra.
The
absence
of
the
bluish
discoloration
is
diagnostic
for
L.
scabrum
and
L.
rotundifoliae.
However,
the
bluish
discoloration
is
not
always
clearly
observable
and
should
therefore
be used
with
caution.
Other
colour
changes
of
the
context,
such
as
yellow
and
red
discolorations
in
the
stipe
base
and
an
olivaceous
discoloration
in
the
apex
of
the
stipe
are
variable
within
species.
In
particular
the
olivaceous
discoloration
in
the
apex
of
the
stipe
seems
to
be
associated
either
with
a
wet
growing
habitat
or
continuous
wet
weather.
For
this
reason
they
are
considered
of
very
limited
diagnostic
value.
Macrochemical
reactions
Lannoy
&
Estades
(1995)
considered
the
reaction
of
the
context
with
FeS0
4
and
Formol
to
be
of
importance
for
the
identification
of
certain
species
groups.
When
FeS04
crystals
are
rubbed
against
the
context
of
(preferably)
the
apex
of
the
stipe,
a
green-
ish
grey
to
blackish
discoloration
may
appear.
Formol,
when
applied
to
the
context,
sometimes
induces
a
pinkish
to
reddish
discoloration
of
the
context.
The
usefulness
of
FeS0
4
as
a
reagent
for
the
identification
of
species
in
subsection
Scabra
has
been
tested.
It
was
often
found
to
be
very
gradual,
and
difficult
to
assess
whether
the
discoloration
was
greyish
green
or
greyish.
Moreover,
there
seems
to
be
a
relation
between
age
and
humidity
of
the
fruit-body
and
the
intensity
of
the
reaction
with
either
formaldehyde
or
FeS0
4
.
Therefore
these
macrochemical
reactions
have
not
been used
in
the
present
work.

526
PERSOONIA
-
Vol.
18,
Part
4,
2005
Colour
of
pileus
and
overall
colour
of
fruit-body
The
colour
of
the
pileus
has
always
been
an
important
diagnostic
character
in
Lec-
cinum
taxonomy.
In
particular
new
species
have been
based
on
an
overall
pale
appear-
ance
or
very
pale
pileus
colour.
Often
authors
considered
these
species
as
pale-coloured
sister
species
to
darker
coloured
species.
Lannoy
&
Estades
(1995)
suggested
that
L.
cyaneobasileucum
is
a
pale
form
of
L.
brunneogriseolum,
by
indicating
that
it
resem-
bles
the
latter
species
except
for
the
overall
colour
of
the
fruit-bodies.
The
molecular
data
(Den
Bakker
et
al.,
2004b,
2005)
suggest
that
most
species
display
a
whole
colour
range
from
light
to
dark.
Based
on
these
findings
it
can
be
concluded
that
for
instance
L.
cyaneobasileucum
is
a
pale
variant
ofL.
brunneogriseolum,
and
that
L.
avellaneum
and
L.
roseofractum
represent
the
lightest
and
darkest
limits
respectively
of
L.
scabrum.
Also
L.
roseotinctum
and
L.
percandidum
are
just
pale-coloured
forms
of
L.
versipelle.
In
addition,
molecular
data
show
that
the
generally
white
or
light-coloured
species
L.
holo-
pus
may
form
fruit-bodies
with
a
brown
pileus
and
dark
squamules
on
the
stipe.
Within
subsection
Leccinum,
however,
there
is
a
clear
distinction
between
species
with
a
dark
reddish
pileus
colour
that
changes
to
dark
reddish
brown
in
exsiccates
(like
L.
aurantiacum
and
L.
vulpinum
?)
and
species
with
a
orange-brown
pileus
colour
that
changes
to
light
brown
in
exsiccates
(like
L.
versipelle
and
L.
albostipitatum).
Stipe
squamules
and
stipe
surface
colour
While
the
colour
of
the
pileus
is
variable
in
most
species,
the
colour
of
the
stipe
surface
can
be
useful
in
the
identification
of
Leccinum
species.
Care
should
be
taken
with
overall
pale-coloured
fruit-bodies
and
fruit-bodies
that
are
found
in
high
and
dense
vegetation.
In
the
first
case
fruit-bodies
might
be
from
a
mycelium
that
produces
a
limited
amount
of
dark
pigments,
in
the
second
case
shade
can
prevent
the
formation
of
dark
pigments
(personal
observation
of
the
first
author).
In
subsection
Leccinum
a
distinction
can
be
made
between
species
with
a
basically
greyish
to
blackish
stipe
ornamentation
(L.
vulpinum
and
L.
versipelle)
and
species
with
a
whitish
to
reddish
brown
stipe
ornamentation
(L.
aurantiacum
and
L.
albostipitatum).
In
subsection
Scabra
most
species
have
a
greyish
to
blackish
stipe
ornamentation,
with
the
exception
of
L.
holopus
var.
holopus,
which
has
an
initially
white
stipe
ornamenta-
tion
that
becomes
darker
in
mature
fruit-bodies,
and
L.
cyaneobasileucum,
which
has
an
initially
whitish
stipe
ornamentation
that
becomes
dirty
greyish
in
older
fruit-bodies.
The
shape
and
pattern
of
the
stipe
squamules
is
of
diagnostic
value
as
well,
especially
in
subsection
Scabra.
The
stipe
squamules
are
usually
fine
at
the
apex
and
coarser
at
the
base
of
the
stipe.
Leccinum
scabrum
and
L.
holopus
usually
have
overall
fine
squamules
on
the
stipe,
whereas
L.
cyaneobasileucum
usually
has
a
stipe
covered
with
coarse,
flocculose
squamules.
While
in
most
species
the
squamules
are
more
or
less
conical,
some
species
(like
L.
rotundifoliae)
have
more
warty
squamules.
The
colour
of
the
stipe
surface
beneath
the
squamules
is
white
or
whitish
in
most
European
Leccinum
species
with
the
exception
of
L.
crocipodium
with
yellow
back-
ground
colour,
and
L.
melaneum
in
which
the
surface
is
distinctly
greyish.
Pileus
surface
The
pileus
surface
in
species
of
subsections
Leccinum,
Fumosa
and
Scabra
is
felted
to
fibrillose.
In
subsection
Leccinum
the
fibrils
on
the
pileus
surface
may
develop
into

527
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
a
pattern
of
appressed
squamules.
The
surface
may
become
viscid
or
slightly
slimy
in
mature
and
wet
fruit-bodies.
The
surface
of
L.
pseudoscabrum
is
often
distinctly
rugulose,
especially
when
young,
whereas
that
of
L.
crocipodium
is
tomentose.
Both
in
L.
pseudoscabrum
and
L.
crocipodium
the
surface
of
the
pileus
usually
cracks
with
age,
especially
during dry
periods.
Other
species
only
show
a
cracked
surface
under
extremely
dry
circumstances.
Pileus
margin
Most
species
of
Leccinum
have
an
entire
margin
that,
especially
in
younger
fruit-
bodies,
projects
a
few
millimeters
beyond
the tubes.
In
subsection
Leccinum
the
margin
of
the
pileus
is
inflexed
in
young
fruit-bodies,
becoming
disrupted
and
appendiculate
when
the
fruit-body
ages.
ECOLOGICAL
CHARACTERS
Host
specificity
Host
specificity
is
often
considered
an
important
character
for
taxonomy
(see
Den
Bakker
et
al.,
2004b
for
further
references).
Species
of
subsection
Scabra
are
associ-
ated
with
Betula,
while
species
of
subsection
Fumosa
are
associated
with
Populus.
Leccinum
montanum
i
l9
and
L.
californicum
20
are
two
species
of
the
Sierra
Nevada
(California,
USA)
associated
with
Populus
(Thiers,
1975).
Based
on
the
absence
of
a
blackish
discoloration
of
the
context
and
their
entire
pileus
margin
Thiers
referred
them
to
subsection
Scabra.
The
habitus
of
these
species
and
their
association
with
Populus
suggests
that
they
are
actually
more
closely
related
to
species
of
subsection
Fumosa.
In
subsection
Leccinum
species
are
found
that
are
exclusively
associated
with
Populus
(L.
albostipitatum),
with
Betula
(L.
versipelle
and
L.
atrostipitatum
21
),
with
Pinaceae
(L.
vulpinum,
L.
piceinum)
)
and
with
Ericaceae
that
form
arbutoid
mycorrhizas
(L.
man-
zanitae
22
and
L.
monticola
2i
).
In
Europe
the
only
species
that
is
not
host
specific
is
L.
aurantiacum.
This
species
is
associated
with
Quercus
and
other
Fagaceae,
Populus
and
Salix,
Betula
and
sometimes
with
Tilia.
Although
many
species
within
Leccinum
are
host
specific,
the
use
of
host
associations
in
the
field
as a
diagnostic
character
is
often
difficult,
since
usually
more
than
one
host
is
present
in
the
vicinity
of
a
fruit-body.
Edaphic
factors
Especially
in
subsection
Scabra
the
acidity
and
the
humidity
of
the
soil
appear
to
be
im-
portant
factors
determining
the
distribution
of
the
different
species.
While
most
species
of
subsection
Scabra
are
found
in
acidic
habitats,
L.
schistophilum
is
found
in
basic
habitats.
Leccinium
holopus
var.
holopus
is
found
in
waterlogged
Sphagnum
vegetation,
L.
holo-
pus
var.
americanum
is
found
in
less
wet,
marshy
Betula
forests.
Also
in
wet
but
not
waterlogged
habitats
species
like
L.
schistophilum,
L.
cyaneobasileucum,
and
L.
varii-
color
can
be
found,
whereas
L.
scabrum
seems
to
be
a
species
of
drier
habitats.
19)
Leccinum
montanum
Thiers,
Mycologia
63
(1971)
274.
20)
Leccinum
californicum
Thiers,
Mycologia
63
(1971)
273.
21)
Leccinum
atrostipitatum
A.H.
Sm„
Thiers
&
Watling,
Michigan
Bot.
5
(1966)
555.
22)
Leccinum
manzanitae
Thiers,
Mycologia
63
(1971)
226.
23)
Leccinum
monticola
Hailing
&
G.M.
Muell.,
Mycologia
95
(2003)
493.

528
PERSOONIA
-
Vol.
18,
Part
4,
2005
MICROSCOPICAL
CHARACTERS
Anatomy
of
pileipellis
The
anatomy
of
the
pileipellis
has
received
much
attention
in
the
taxonomy
of
Leccinum
(Blum,
1970;
Smith
&
Thiers,
1971;
Lannoy
&
Estades,
1995).
Within
the
genus
Leccinum,
two
basic
types
can
be
recognized,
viz.
(1)
a
trichoderm
of
erect
chains
of
elements,
sometimes
forming
a
palisade,
typical
for
L.
pseudoscabrum
and
L.
crocipodium
(see
Fig.
4A)
and
(2)
a
complex
type
of
intricate
trichoderm,
often
with
a
cutis-like
suprapellis,
typical
for
subsections
Leccinum,
Fumosa
and
Scabra
(Figs.
4
C,
D
and
E).
Although
this
cannot
be
considered
a
real
cutis,
and
in
young
fruit-bod-
ies
of
species
of
subsection
Scabra
sometimes
a
real
trichoderm
without
a
cutis-like
suprapellis
can
be
observed,
we
will
for
convenience
call
this
type
of
pileipellis
a
cutis
in
the
species
descriptions.
Particular
in
subsection
Scabra
pileipellis
structures
appear
to
be
rather
variable.
Lannoy
&
Estades
(1995)
considered
the
presence
or
absence
of
so-called
cylindrocysts
in
the
pileipellis
an
important
character
for
certain
species
in
subsection
Scabra.
They
defined
cylindrocysts
as
hyphal
elements
that
are
maximal
80
pm
long
and
minimal
10
pm
wide
and
have
length/width
ratio
of
less
than
4.
According
to
Lannoy
&
Estades
the
presence
of
cylindrocysts
is
diagnostic
for
their
subsection
Pseudoscabra.
The
present
study
revealed
that
the
occurrence
of
cylindrocysts
is
not
consistently
correlated
with
the
current
species
concept.
Furthermore,
no
indication
was
found
that
Fig.
4.
Overview
of
pileipellis
types
encountered
in
Leccinum
(radial
sections,
scale
bars
=
10
µm).
A.
A
trichoderm,
as
found
in
B
to
E.
cutis-like
structures,
named
cutis
in
the
species
descriptions
(B.
L.
crocipodium;
L.
variicolor,
C.
L.
versipelle,
L.
holopus
The
pileipellis
of
D.
and
E.
L.
scabrum).
L.
variicolor,
as
depicted
here,
is
mainly
composed
of
so-called
‘cylindrocysts’.

529
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
subsection
Pseudoscabra
forms
a
natural
group.
Species
with
and
without
cylindrocysts
have been
synonymised (L.
rigidipes
and
L.
scabrum,
L.
aerugineum
and
L.
holopus).
However,
in
some
species,
like
L.
variicolor
and
L.
cyaneobasileucum
cylindrocysts
are
usually
abundant,
and
may
facilitate
identification.
Pigmentation
of
pileipellis
The
pigmentation
of
the
pileipellis
elements
can
be
granularly
incrusted,
plaque-like
incrusted
or
vacuolar.
The
pigmentation
type
varies
within
species.
Certain
pigmenta-
tion
types
occur
more
frequently
in
certain
species.
For
instance
the
pileipellis
elements
of
L.
schistophilum
and
L.
aurantiacum
are
often
plaque-like
incrusted.
The
colour
of
the
pigments
observed
usually
corresponds
to
the
overall
colour
of
the
pileus.
Smith
et
al.
(1967)
and
Watling
(1970)
introduced
another
character
for
discriminating
species
in
subsection
Leccinum,
viz.
the
capacity
of
the
pigments
to
form
globules
when
the
pileipellis
is
mounted
in
Melzer's
reagent.
This
reaction
did
not
appear
to
be
constant
within
species,
and
could
not
be used
as
diagnostic
character.
Spore
size
and
spore
shape
Spore
shape
appeared
to
be
a
better
diagnostic
character
than spore
size
in
discrimi-
nating
species.
In
subsection
Leccinum
the
mean
Q-value
(length
:
width
ratio)
varies
between
3.1
and
3.5,
and
in
subsection
Scabra
between
2.7
and
2.8,
with
the
exception
of
the
European
collections
of
L.
scabrum
and
L.
cyaneobasileucum
in
which
a
mean
Q-value
of
3.0
and
3.2
was
observed.
In
the
collections
of
L.
scabrum
from
eastern
North
America
the
mean
Q-value
was
2.8.
Since
the
mean
Q-value
was
observed
to
be
constant
in
the
species
of
subsection
Scabra
in
Europe,
this
could
be used
as
an
ad-
ditional
character.
Hymenocystidia
Hymenocystidia
in
the
tubular
hymenium
(pleurocystidia)
are
usually
lageniform.
The
hymenocystidia
found
on
the
pore
surface
(cheilocystidia)
have
various
shapes,
being
either
clavate,
utriform
or
lageniform.
In
subsection
Leccinum
the
cheilocystidia
usually
have
a
brownish
content.
Size
and
shape
of
hymenical
cystidia
were
found
to
be
of
no
diagnostic
value.
Caulocystidia
Thus
far,
caulocystidia
have
remained
relatively
unstudied
in
Leccinum
systematics.
Smith
&
Thiers
(1971)
considered
caulocystidia
too
variable
to
be
of
major
taxonomic
importance,
with
the
exception
of
a
certain
type
of
caulocystidia
in
L.
snellii.
Smith
&
Thiers
considered
the
presence
of
lageniform
cystidia
with
a
septum
(secondary
crosswall)
on
the
transition
between
the
swollen
part
of
the
cystidium
and
the
neck
diagnostic
for
this
species.
Examination
of
other
species
of
subsection
Scabra
showed
that
in
most
species
of
this
subsection
this
septate-lageniform
type
of
caulocystidia
can
be
found,
but
always
in
small
numbers.
In
L.
snellii
and
in
L.
variicolor,
,
these
septate-
lageniform
cystidia
are
usually
common
and
easy
to
observe.
This
is
a
pattern
that
is
seen
repeatedly,
as
in
most
species
in
subsections
Leccinum
and
Scabra
various
types
of
caulocystidia
can
be
found,
but
in
some
species
certain
types
seem
to
be
more
numerous
than
other
types.
As
such
caulocystidia
can
therefore
be used
as
an
additional
character
in
the
identification
of
individual
species.

PHRSOONIA
-
Vol.
18,
Part
4,
2005530
TAXONOMIC
PART
LECCINUM
GRAY
Leccinum
Gray,
Nat.
Arr.
Brit.
PI.
1(1821)646.
-J
Krombholzia
P.
Karst.,
Rev.
Mycol.
(Paris)
3
(1881)
17,
non
Krombholzia
Rupr.
ex
E.
Fourn.
1876;
Krombholziella
Maire,
Publ.
Inst.
Bot.
3
(4)
(1935)
41.
-
Trachypus
Bataille,
Bolets
(1908)
12,
non
Trachypus
Reinw.
&
Hornsch.
1829.
—
Leccinellum
Bresinsky
&
Binder,
Regensb.
Mykol.
Schr.
11
(2003)231.
Selected
literature.
Watling,
Br.
Fung.
Fl.
1
(1970)
45-60;
Lannoy
&
Estades,
Mon.
Leccinum
(1995);
Estades
&
Lannoy,
Bull.
Trimestriel
Fed.
Mycol.
Daupine-Savoie.
174
(2004)60-73.
Basidiocarp
boletoid;
pileus
fibrillose,
tomentose
or
rugulose
and
smooth;
tube
layer
subventricose
to
broadly
ventricose,
adnexed
to
narrowly
adnate;
pores
roundish,
0.3
-
0.5
mm
in
diameter,
whitish
or
yellowish.
Stipe
squamulose.
Context
whitish,
in
some
species
yellowish,
discolouring
greyish,
bluish
or
pinkish
when
bruised,
some-
times
not
discolouring.
Spores
fusiform
to
broadly
fusiform,
smooth;
cheilo-
and
pleurocystidia
present;
pileipellis
an
(ixo-)cutis
or
trichoderm;
pigment
in
pileipellis
intracellular,
vacuolar,
incrusting
or
parietal;
clamp-connections
absent.
—
Type
species:
Leccinum
aurantia-
cum
(Bull.)
Gray.
Habitat
&
distribution
—
Ectomycorrhizal.
Northern
Hemisphere.
Note.
In
the
nomenclator
accompanying
species
descriptions,
the
combinations
in
Krombholzia
and
Krombholziella
have
not
been
listed.
KEY
TO
WESTERN
EUROPEAN
SPECIES
OF
LECCINUM
la.
Context
almost
instantly
discolouring
greyish
to
blackish
when
bruised
2
b.
Context
not
discolouring
instantly
greyish
blackish
when
bruised,
sometimes
dis-
colouring
greyish
after
several
hours
9
2a.
Hymenium
yellowish
13.
L.
crocipodium
b.
Hymenium
whitish,
greyish
or
brownish
3
3a.
Margin
of
the
pileus,
especially
in
young
fruit-bodies,
overhanging
with
disrupted
flaps,
colour
of
pileus
usually
reddish
brown
or
orange,
sometimes
pinkish
or
whit-
ish
4
b.
Margin
of
the
pileus
entire,
only
slightly
projecting;
colour
of
pileus
brownish
.
8
4a.
Stipe
ornamentation
whitish,
brownish
to
reddish
brown,
in
older
specimens
some-
times
almost
blackish;
content
of
caulocystidia
reddish
brown;
associated
with
broadleaved
trees
5
b.
Stipe
ornamentation
brown
or
black,
content
of
pigmented
caulocystidia
greyish,
associated
with
Pinaceae
or
Betula
6
5a.
Stipe
in
young
fruit-bodies
with
reddish
brown
squamules,
which
becomedarker
with
age;
pileus
reddish
brown,
dark
(reddish)
brown
in
exsiccates
1.
L.
aurantiacum

531
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
b.
Stipe
in
young
fruit-bodies
with
whitish
squamules,
which
becomedarker
only
in
very
mature
stage;
pileus
orange,
light
brown
in
exsiccates
2.L.
albostipitatum
6a.
Pileus
colour
usually
orange
or
brownish
orange,
largest
caulocystidia
generally
utriform,
fusiform,
sometimes
lageniform,
associated
with
Betula
4.L.
versipelle
b.
Pileus
brownish
to
reddish
brown,
largest
caulocystidia
generally
clavate,
associ-
ated
with
Pinaceae
7
7a.
Pileusreddish
brown,
associated
with
Pinus
or
Picea
3.
L.
vulpinum
b.
Pileus
dull
brownish,
associated
with
Picea
L.
piceinum
(see
discussion
L.
vulpinum)
8a.
Surface
of
pileus
wrinkled
in
young
fruit-bodies;
pileipellis
a
palisade
trichoderm
of
broadly
clavate
elements,
terminal
elements
often
globose;
associated
with
Corylus
or
Carpinus
4. L.
pseudoscabrum
b.
Surface
of
pileus
never
wrinkled;
pileipellis
a
cutis
with
ellipsoid
or
filiform
termi-
nal
elements;
associated
with
Populus
5.
L.
duriusculum
9a.
Cylindrocysts
in
pileipellis
abundant;
stipe
base
usually
with
clear
greenish
bluish
stains
10
b.
Cylindrocysts
in
pileipellis
scarce;
stipe
base
with
or
without
bluish
stains
...
11
10a.
Stipe
squamules
coarse
and
usually
concolorous
to
stipe
surface,
forming
a
fibril-
lose
dense
layer;
pileus
evenly
coloured;
pileipellis
elements
usually
easily
detach-
able,
often
broad
(8-15
pm);
mean
Q-value
spores
>
3.0
1
\
.
L.
cyaneobasileucum
b.
Stipe
squamules
fine,
usually
blackish,
greyish
and
contrasting
to
the
stipe
surface;
pileus
often
dark-coloured
with
lighter
regions
or
light-coloured
with
darker
re-
gions;
pileipellis
consisting
of
an
intricate
cutis
of
filamentous
hyphae,
often
with
chains
of
cylindrocysts
in
the
suprapellis;
mean
Q-value
spores
<
2.8
10.
L.
variicolor
11a.
Stipe
surface
dark
(greyish,
blackish)
almost
concolorous
to
stipe
squamules
7.
L.
melaneum
b.
Stipe
surface
light,
brownish,
strongly
contrasting
to
squamules
when
squamules
are
darkly
coloured,
otherwise
squamules
light
and
not
contrasting
to
the
surface
12
12a.
Mean
Q-value
spores
s=3.0,
usually
large
clavate
caulocystidia
present
in
caulohy-
menium,
bluish
discolorations
absent
in
stipe
base
6. L.
scabrum
b.
Mean
Q-value
spores
<
3.0,
no
large
clavate
caulocystidia
present
in
caulohy-
menium,
bluish
stainsabsent
or
present
in
stipe
base
13
13a.
Caulocystidia
often
clavate,
sometimes
lageniform;
bluish
discoloration
in
stipe
base
absent
8.
L.
rotundifoliae
b.
Caulocystidia
narrowly
lageniform
with
a
flexuose
neck
or
utriform,
rarely
clavate;
bluish
discoloration
in
stipe
base
present
or
absent
14
14a.
Stipe
ornamentation
composed
of
coarse
warty
or
flocculose
squamules;
caulo-
cystidia
often
narrowly
lageniform
with
a
flexuose
neck;
pileipellis
elements
with
a
vacuolar
pigment
or
granualar
incrustation;
pileus
usually
whitish,
light
brownish,
sometimes
with
bluish
or
greenish
tinges
12.
L.
holopus

PERSOONIA
-
Vol.
18,
Part
4,
2005532
b.
Stipe
ornamentation
composed
of
fine
greyish
or
blackish
squamules;
caulocystidia
often
utriform,
rarely
lageniform
with
a
flexuose
neck;
pileipellis
elements
often
incrusted
with
dark
plaques
of
pigment;
pileus
greyish
or
brownish
9.L.
schistophilum
SECTION
LECCINUM
SUBSECTION
LECCINUM
Pileal
margin,
especially
in
young
fruit-bodies,
with
appendiculate
flaps.
Context
whitish,
showing
bluish,
pinkish,
reddish
and
greyish/blackish
discolorations
when
bruised.
Widely
distributed
throughout
the
Northern
Hemisphere.
Type
species:
Leccinum
aurantiacum
(Bull.)
Gray.
1.
Leccinum
aurantiacum
(Bull.)
Gray
—
Fig.
5,
Plate
1
Boletus
aurantiacus
Bull.,
Herb.
Fr.
(1785)
pi.
236;
Leccinum
aurantiacum
(Bull.)
Gray,
Nat.
Arr.
Br.
PI.
1
(1821)646.
—
Boletus
leucopodius
Pers.,
Observ.
My
col.
2
(1799)
11;
Boletus
aurantiacus
6
[var.]
leucopodius
(Pers.)
Pers.,
Syn.
Meth.
Fung.
(1821)
504;
Leccinum
aurantiacum
B
[var.]
leu-
copodium
(Pers.)
Gray.
Nat.
Arr.
Brit.
PI.
(1821)
646;
Leccinum
leucopodium
(Pers.)
Dorfelt
&
G.
Berg,
Fedd.
Repert.
101 (1990)
567.
—
Leccinum
aurantiacum
var.
quercinum
Pilat,
Mushr.
Other
Fungi
(1961)
6
(footnote)
[invalid,
no
Latin
diagnosis];
Leccinum
quercinum
(Pilfit)
Green
&
Watling,
Notes
Roy.
Bot.
Gard.
Edinburgh
29
(1969)
265
[invalid,
basionym
not
validly
published];
Leccinum
quercinum
Pilat
in
Pilat
&
Dermek,
Hrlbovite
huby
(1974)
151.
—
Leccinum
salicola
Watling,
Notes
Roy.
Bot.
Gard.
Edinburgh
31
(1971)
139.
—
Leccinum
populinum
M.
Korhonen,
Karstenia35
(1995)
55.
Iconotype.
Bulliard
1785.
Herb.
Fr. PI.
236.
Epitype
(designated
here).
France:
Orne,
Foret
de
Belleme,
28IX
2001,
J.
van
Brummelen
(nr.
11
+
12)(L).
Misapplication.
Leccinum
rufum
sensu
auct.
Excluded.
L.
aurantiacum
sensu
M.
Korhonen,
Karstenia
35
(1995)
58
(=
L.
albostipitatum);
L.
leucopodium
sensu
Den
Bakker
et
al.,
New
Phytol.
163 (2004)
201-215
(=
L.
albostipitatum).
Selected
icones.
Lannoy
&
Estades,
Mon.
Lecc.
Europe
(1995)
pi.
37
(as
L.
quercinum)
;
Korhonen,
Karstenia
35
(1995)
54,
fig.
1
&
2
(as
L.
populinum
and
L.
quercinum
respectively).
Selected
descriptions.
Korhonen,
Karstenia
35
(1995)
55-58
(as
L.
populinum
and
L.
quercinum
respectively).
Characteristics
—
A
slender
to
robust,
medium
to
large
Leccinum;
pileus
reddish
brown,
dark
reddish
brown
in
exsiccates;
stipital
squamules
reddish
in
young
fruit-
bodies,
dark
reddish
brown
in
older
fruit-bodies,
seldom
whitish.
Mycorrhizal
with
various
kinds
of
broad-leaved
trees
(
Populus,
Salix,
Quercus,
Fagus,
Betula,
Castanea,
Tilia).
Pileus
30-150
(-280)
mm,
truncate-conical
to
conico-convex
or
hemispherical
when
young,
expanding
to
truncate
conico-convex,
convex
or
plano-convex
when
mature,
with
involute
then
deflexed
margin,
extending
over
tubes
with
distinct
triangular
or
irregularly
shaped
flaps,
up
to
6
mm
long,
very
evident
in
young
specimens,
and
generally
also
in
mature
specimens,
rarely
disappearing
with
age;
vivid
red
to
red-brown
(Mu.
2.5
YR
4/6
-5/8;
10
R
3-4/3-6),
towards
margin
sometimes
more
orange-yellow
(7.5
YR
7/8),

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
533
Leccinum
aurantiacum.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.
Fig.
5.

PERSOONIA
-
Vol.
18,
Part
4,
2005534
initially
finely
tomentose,
breaking
up
with
age
in
small
appressed
squamules
all
over.
Tubes
adnexed
to
adnate,
segmentiform
to
ventricose,
9-30
mm
long,
cream-white
to
very
pale
brown
(10
YR
6/4).
Pores
c.
0.2-0.5
mm
in
diameter,
yellowish
white
to
very
pale
brown
(10
YR
7/4),
discolouring
brownish
when
bruised.
Stipe
50-270
x
15-50
mm,
cylindrical
to
clavate
to
fusiform,
whitish,
often
with
a
distinct
blue
discoloration
at
base,
especially
when
handled,
entirely
covered
with
squamules,
which
are
small
and
isolated
at
apex,
and
coarse
and
more
dense
and
sometimes
agglutinated
towards
base,
initially
reddish,
darkening
to
red-brown
or
reddish
black
with
age.
Context
white,
turning
violaceous
grey,
grey
or
blackish
when
bruised;
in
stipe
base
often
turning
blue-
green.
Spores
12.5-18.5
(-22.5)
x
3.5-5.0(-6.0)
pm,
Q
=
2.8
-4.1,
Q
av
=
3.2-3.6;
fusiform
with
conical
apex,
with
distinct
suprahilar
depression,
pale,
purplish
brown
in
water.
Basidia
15
-25.0
x
6.5-10.5
pm,
clavate,
2-
or
4-spored.
Hymenocystidia
20-45
x
5-8
pm,
narrowly
lageniform
and
hyaline
in
tubular
hymenium,
clavate
or
utriform
with
reddish
brown
content
on
pore
surface.
Pileipellis
an
intricate
trichoderm
of
(narrowly)
cylindrical
elements,
terminal
elements
often
conical
or
cylindrical
with
a
mucronate
apex,
30-100
x
3.0-15.5
p
m,
elements
in
suprapellis
50
-
70
x
5.5-12.5
pm,
pigment
in
water
vacuolar
or
in
globules,
larger
elements
often
intracellularly
granular
incrusted,
red-brown.
Caulocystidia
30-
60(-90)
x
(6.5-)
12.5-20.5
pm,
clavate
with
acute
or
obtuse
apex
to
almost
spheropedunculate,
rarely
utriform
or
fusiform,
often
in
addition
small
lageniform
cystidia
present;
large
cystidia
usually
with
reddish
brown
content
in
KOH;
small
lageniform
cystidia
usually
hyaline.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Popu-
lus,
Quercus
and
Betula,
rarely
with
Salix,
Fagus,
Castanea
and
Tilia,
on
mesotrophic,
sandy
or
loamy
soils;
moderately
common
in
the
Netherlands,
widespread
in
Europe.
Collections
studied.
THE
NETHERLANDS:
Drenthe,
Gieten,
Boekweitveen,
16
X
2000,
H.C.
den
Bakker
54:
Roden,
20
IX
2000,
J.
Wisman,
H.C.
den
Bakker
102;
Gelderland,
Heiligenberg,
2001,
J.
Wisman,
H.C.
den
Bakker
286
;
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
14
VIII
1999,
H.C.
den
Bakker
3.
—
BELGIUM:
Namur,
Oignies-en-Thierache,
22
IX
1999,
Ruben
Walleyn
1683.
—
CZECH
REPUBLIC:
Bohemia,
distr.
Turnov,
Masov-Sedmihorky,
H.
Schmidt,
PRM521031
(holotype
L.
quercinum,
PRM)-ENGLAND:
Berkshire,
Windsor
Great
Park,
2001,4.
Hills
2001219.
—
FINLAND:
Nylandia,
Vantaa,
12
IX
1994,
M.
Korhonen
11850
(holotype
L.
populinum,
H).
—
FRANCE:
Orne,
Foret
de
Belleme,
28
IX
2001,
J.
van
Brummelen
(Fix.
3
+
4);
Orne,
Foret
de
Belleme,
28
IX
2001,
J.
van
Brummelen
(Fix.
11
+
12);
Orne,
Ige,
3
X
2001,7.
van
Brummelen
(Fix.
17
+
18);
Doubs,
ForSt
de
Cessey,
8
X
2001,7.
van
Brummelen
(Fix.
19+20).
—
NORWAY:
Sogn
og
Fjordane,
Sogndal,
7IX
2000,
H.C.
den
Bakker
94.
—
SCOTLAND:
Invemess-shire,
Insh
Nature
Reserve,
17IX
2001,
H.C.
den
Bakker
233;
Sutherland,
Glaisgeo,
22IX
1970,
Watling
7362
(holotype
L.
salicola,
E).
Notwithstanding
intensive
molecular
and
morphological
studies
of
collections
(in-
cluding
holotypes)
that
were
associated
with
Populus,
Quercus
and
Betula,
no
clear
support
could
be
found
for
the
recognition
of
L.
populinum
and
L.
quercinum
as
distinct
species.
Therefore
we
merge
these
taxa
into
one
species,
L.
aurantiacum,
with
a
broad
host
range
(Den
Bakker
et
al.,
2004b).
No
obvious
differences
in
spore
size
or
in
spore
dimensions
were
found
between
collections
that
were
associated
with
either
Populus
or
Quercus
(Fig.
6),
in
contrast
to
the
observations
by
Korhonen
(1995).
Kibby
(2002)
in-
dicated
that
the
shape
of
the
caulocystidia
of
L.
populinum
differs
from
those
of
L.
quer-
cinum
Leccinum
populinum
is
supposed
to
have
inflated,
clavate
caulocystidia,
while
the
caulocystidia
are
usually
fusiform
capitate
in
L.
quercinum.
The
present
study
revealed

535
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
that
caulocystidia
as
indicated
by
Kibby
for
L.
populinum
can
be
found
in
most
collec-
tions
of
L.
populinum
and
L.
quercinum
(including
the
holotype
of
L.
quercinum)
and
that
fusiform
capitate
caulocystidia
are
more
rarely
encountered
in
Populus
or
Quercus
associated
collections.
Leccinum
salicola
Watling
was
described
by
Watling
(1971)
and,
according
to
the
author,
could
be
..
distinguished
from
all
other
members
of
the
L.
aurantiacum
com-
plex
by
the
cinnamon
to
brick-coloured
pileus,
coupled
with
the
habitat
and
(small
and
stocky)
stature
and
the
(pallid
or
sepia)
pore
colour
when
young..The
small
stocky
stature
of
this
taxon
is
probably
a
result
of
the
exposed
nature
of
its
habitat
(
Salix
re-
pens
vegetation
on
cliff-tops),
and
both
pileus
and
pore
colour
do
not
differ
from
our
concept
of
L.
aurantiacum.
A
study
of
the
type
material
showed
that
Leccinum
salicola
fits
well
into
our
concept
of
L.
aurantiacum.
Leccinum
salicola
is
therefore
considered
a
synonym
of
L.
aurantiacum.
Several
authors
(Dorfelt
&
Berg
1990;
Redeuilh,
1990;
Sutara,
1989)
have
discussed
the
identity
of
Bulliard's
type
plate
of
L.
aurantiacum
and
came
to
the
conclusion
that
it
represents
L.
quercinum.
Our
results
indicate
that
there
is
no
sound
morphological
basis
(∆),
Quercus
(o),
associated
with
Populus
Fig.
6.
Spore
size
in
basidiocarps
ofof
L.
aurantiacum
Salix
Betula
(□).
(x)
and

PERSOONIA
-
Vol.
18,
Part
4,
2005536
for
considering
L.
quercinum
a
distinct
species
and
we
therefore
conclude
that
Bulliard
depicted
the
same
taxon
we
currently
call
L.
aurantiacum.
In
Den
Bakker
et
al.
(2004b)
we
followed
the
recommendations
of
Redeuilh
(1990)
and
called
L.
aurantiacum
sensu
Korhonen
L.
leucopodium
(Pers.)
Dorfelt
&
G.
Berg.
Careful
examination
of
the
study
of
Dorfelt
&
Berg
(1990),
however,
showed
that
their
interpretation
ofL.
leucopodium
is
the
same
as
our
concept
of
L.
aurantiacum.
They
had
introduced
the
name
L.
leuco-
podium
for
the
taxon
growing
under
Populus,
as
the
name
L.
aurantiacum
was
applied
for
the
taxon
associated
with
Quercus.
Therefore
L.
aurantiacum
sensu
Korhonen
is
described
as a
new
species
(L.
albostipitatum),
in
order
to
avoid
taxonomic
confusion
about
this
taxon.
2.
Leccinum
albostipitatum
H.C.
den
Bakker
&
Noordel.,
spec.
nov.
—
Fig.
7,
Plate
2
Leccino
aurantiaco
similis,
pileo
aurantiaco,
stipite
squamulis
albis
demum
parum
rubiginosis.
Populo
consociatus.
Holotypus:
Italy,
Trento,
Monte
Bella,
10
IX
1996.
M.E.
Noordeloos
96134
(L).
Misapplication.
Leccinum
aurantiacum
sensu
auct
p.p.;
sensu
Korhonen,
Karstenia
35
(1995)
58.
—
Leccinum
leucopodium
Pers.
sensu
Den
Bakker,
New
Phytol.
163 (2004)
201-215.
Selected
icones.
Lannoy
&
Estades,
Mon.
Leccinum
(1995)
pi.
34
(as
L.
aurantiacum)
);
Korhonen,
Karstenia
35
(1995)
55,
fig.
3
(as
L.
aurantiacum).
Selected
description.
Korhonen,
Karstenia
35
(1995)
58-61
(as
L.
aurantiacum).
Characteristics
—
Habit
like
L.
aurantiacum;
pileus
light
orange,
light
brown
in
exsiccates;
stipital
squamules
whitish
in
young
fruit-bodies,
in
older
fruit-bodies
light
brownish.
Mycorrhizal
with
Populus.
Pileus
80-
250
mm,
hemispherical
when
young,
convex
to
plano-convex
when
mature;
margin
especially
in
young
fruit-bodies
inflexed,
with
appendiculate
flaps,
ex-
ceeding
up
to
4
mm,
in
mature
fruit-bodies
margin
often
seemingly
entire
and
marginal
flaps
absent,
vivid
orange
(Mu.
7.5
YR
7/6-7/8);
surface
minutely
tomentose
to
fibrillose
squamulose.
Tubes
adnexed
to
adnate,
ventricose,
9-30
mm
long,
pale
brownish
(10
YR
6/4),
yellowish
white.
Pores
c.
0.5
mm
in
diameter,
yellowish
white
to
very
pale
brownish
(10
YR
7/4),
discolouring
brownish
when
bruised.
Stipe
50-270
x
15—
50
mm,
cylindrical
to
subclavate,
whitish,
often
with
a
clear
blue
discoloration
in
the
stipe
base
when
handled,
sparsely
to
densely
covered
with
fine
(sometimes
on
the
stipe
base
more
coarse),
whitish
(when
young)
to
reddish
brown
(in
older
specimens) squamules.
Context
white,
when
bruised
often
discolouring
first
vinaceous,
than
greyish,
blackish,
often
discolouring
bluish
in
the
stipe
base.
Spores
(9.5-)l
1.0-17.0
x
4.0-5.0(-5.5)
pm,
Q
=
2.3-3.6(-4.0),
Q
av
=
(2.9-)
3.0-
3.1;
fusiform with
a
conical
apex,
with
a
distinct
suprahilar
depression,
pale,
purplish
brown
in
water.
Basidia
25-35
x
7.5-11.0
pm,
clavate,
2-
or
4-spored.
Hymeno-
cystidia
20-45
pm
x
7-10
pm,
lageniform
in
tubular
hymenium,
form
of
cystidia
of
hymenium
pore
surface
not
clearly
observed.
Pileipellis
an
intricate
cutis,
composed
of
narrowly
cylindrical
and
cylindrical
elements,
terminal
elements
often
conical,
35
-
80
x
4.0-15.5
pm,
pigment
in
water
often
granular
incrusted,
yellowish
brown.
Caulocystidia
15-65
x
10.0-16.0
pm,
clavate
to
fusiform,
rarely
lageniform,
largest
cystidia
often
fusiform,
apex
usually
obtuse,
content
pale
sepia
to
hyaline
in
KOH.
Clamp-connections
absent.

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
537
Leccinum
albostipitatum.
Fig.
7.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

538
PERSOONIA
-
Vol.
18,
Part
4,
2005
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Populus
in
mixed
forest
and
Populus
stands;
not
found
in
the
Netherlands,
probably
a
bo-
realand
sub-alpine
species,
common
in
Scandinavia
and
the
mountainous
areas
of
central
Europe,
rare
at
lower
altitudes.
Collections
studied.
FRANCE:
Champagne-Ardenne,
Sommauthe/Beaumont-en-Argonne,
20
IX
1999,
R.
Walleyn
1656.
—
ITALY:
Trento,
Monte
Bella,
10IX
1996,
M.
E.
Noordeloos
96134
(Holo-
type,
L).
—
NORWAY:
Sogn
og
Fjordane,
Sogndal,
7IX
2000,
H.C.
den
Bakker
93.
Leccinum
aurantiacum
has
for
a
long
time
been
broadly
interpreted
and
until
re-
cently
at
least
two
European
species
that
are
associated
with
Populus
were
included
in
the
interpretation
of
this
name.
The
importance
of
the
colour
of
the
stipital
squamules
has
not
been
sufficiently
appreciated.
Korhonen
(1995)
was
the
first
author
to
separate
these
species.
However,
his
proposal
to
use
the
name
L.
aurantiacum
for
the
taxon
with
pale
squamules
can
not
be
accepted
for
various
reasons.
Consequently,
the
recently
de-
scribed
species
L.
populinum
has
to
be
reduced
to
a
synonym
of
L.
aurantiacum,
while
L.
aurantiacum
sensu
Korhonen
is
described
here
as
new,
viz.
L.
albostipitatum.
The
main
difference
between
L.
aurantiacum
and
L.
albostipitatum
is
found
in
the
colour
of
the
stipital
ornamentation
and
the
pileus
colour
both
in
fresh
and
dried
fruit-bodies.
The
first
species
has
generally
a
brownish-reddish
stipital
ornamentation
that
is
already
reddish
in
young
fruit-bodies,
whereas
L.
albostipitatum
starts
with
completely
white
stipital
ornamentation
that
darkens
slightly
in
older
fruit-bodies.
Care
should
be
taken
in
using
this
character
when
the
fruit-bodies
are
found
in
high
vegetation.
We
have
ob-
served
that
the
stipital
ornamentation
of
L.
aurantiacum
can
be
almost
white
when
the
stipe
is
covered
with
grass.
In
these
cases
the
pileus
colour
provides
a
better
character
to
separate
the
two
species.
Leccinum
aurantiacum
has
a
vividly
red
to
reddish
brown
pileus,
and
when
dried
the
pileus
is
reddish
brown.
The
pileus
colour
of
L.
albostipitatum
is
generally
vividly
orange
and
changes
to
a
dull
light
brown
in
dried
fruit-bodies.
Leccinum
aurantiacum
and
L.
albostipitatum
are
the
only
two
European
representa-
tives
of
a
group
(clade)
of
closely
related
species
with
a
(sub-)boreal
distribution.
While
the
species
boundaries
of
the
two
species
encountered
in
Europe
are
clearcut
both
mor-
phologically
and
molecularly
(Den
Bakker
et
al.,
2004b),
the
situation
in
North
America
is
still
taxonomically
confusing
and
in
need
of
a
revision,
which
needs
to
be
supported
with
additional
molecular
data.
Leccinum
aurantiacum
is
probably
a
European
species,
and
no
records
are
known
from
North
America.
The
descriptions
of
L.
aurantiacum
in
North
American
literature
represent
a
mixture
between
a
L.
vulpinum-like,
conifer
associated
taxon
and
North
American
species
that
are
associated
with
broad-leaved
trees,
such
as
L.
insigne,
and
L.
brunneum
24
.
One
L.
aurantiacum-like
collection
from
Ontario,
studied
by
Den
Bakker
et
al.
(2004b),
belonged
to
the
clade
of
L.
albostipitatum.
However,
with
the
current
literature
(Smith
&
Thiers,
1971)
it
was
impossible
to
assign
a
name
to
that
collection.
3.
Leccinum
vulpinum
Watling
—
Fig.
8
Leccinum
vulpinum
Watling,
Trans.
Bot.
Soc.
Edinburgh
39
(1961)
197.
Selected
icones.
Lannoy
&
Estades,
Mon.
Lecc.
Europe
(1995)
pi.
36.
24)
Leccinum
brunneum
Thiers,
Mycologia
63
(1971)
269.

539
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Selected
descriptions.
Watling,
Br.
Fung.
Fl.
1
(1970)
57;
Lannoy
&
Estades,
Mon.
Lecc.
Europe
(1995)138-139.
Characteristics
—
A
slender
to
robust,
medium
to
large
Leccinum;
pileus
reddish
brown,
dark
reddish
brown
in
exsiccates;
stipital
squamules
greyish
to
blackish,
also
in
young
fruit-bodies.
Mycorrhizal
with
Pinaceae.
Pileus
70-92
mm,
hemispherical
when
young,
convex
to
plano-convex
when
ma-
ture,
margin
especially
in
young
fruit-bodies
involute
or
indexed,
with
appendiculate
flaps,
exceeding
up
to
6
mm,
in
mature
fruit-bodies
margin
often
apparently
entire
and
marginal
flaps
absent;
usually
dark
reddish
brown
(Mu.
2.5
YR
6-3/6,
10
R
3/6),
with
patches
of
a
slightly
lighter
colour;
tomentose
to
fibrillose
squamulose.
Tubes
adnate
to
sinuate,
ventricose,
10-16
mm
long,
pinkish
brown
(7.5
YR
7/4)
pale
greyish
brown
(10
YR
7/3-4),
slightly
darker
when
bruised.
Pores
c.
0.5
mm
in
diameter,
concolorous
with
tubes.
Stipe
70-150
x
14-31
mm,
cylindrical
to
subclavate,
sometimes
fusiform,
whitish,
near
the
apex
lightly
covered
with
fine
(sometimes
on
the
stipe
base
more
coarse)
dark
brown
or
greyish
to
blackish
squamules,
towards
the
base
more
densely
covered.
Context
white,
when
bruised
slowly
discolouring
to
greyish
or
blackish
with
a
violet
hue
in
the
pileus
and
upper
half
of
the
stipe,
sometimes
with
bluish
discolorations
in
the
stipe
base.
Spores(9.5
—)
11.0-15.5
x
3.5-4.5
//m,Q=2.5-3.8,
Q
av
=
3.2-3.4(-3.7);
fusiform
with
conical
apex,
with
a
distinct
suprahilar
depression,
pale,
purplish
brown
in
water.
Basidia
20-25
x
7.5-10.5
pm,
clavate.
Hymenocystidia
20-45
x
5-10
pm,
lageniform,
hyaline
in
tubular
hymenium,
pleurocystidia
often
clavate
or
narrowly
utriform
with
a
reddish
brown
content.
Pileipellis
cutis-like,
elements
20-35
x
3.0-3.5
pm,
elements
in
suprapellis
50-70
x
5.5-12.5
pm,
pigment
in
water
vacuolar
or
in
globules,
larger
elements
often
intracellularly
granular
incrusted,
red-brown.
Caulocystidia
20-45
x
9.5-16.0
pm,
generally
clavate
with
an
obtuse
or
acuminate
apex,
content
usually
brownish
in
KOH,
larger
cystidia
usually
hyaline.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
in
clusters,
ectomycorrhizal,
associated
with
Pinus
and
Picea
in
mixed
and
coniferous
forests;
not
known
from
the
Netherlands,
known
from
Scotland,
Scandinavia
and
Central
Europe.
Probably
widespread
in
coniferous
forests
in
subalpine
and
boreal
regions
throughout
the
Northern
Hemisphere.
Collections
studied.
CANADA:
Ontario,
Manitoulin
Island,
30
IX
2003,
H.C.
den
Bakker
415
—
FINLAND:
Vuokatti,
1IX
2001
M.E.
Noordeloos
011243.
—
NORWAY:
Sogn
og
Fjordane,
Sogndal,
7IX
2000,
H.C.
den
Bakker
92.
—
SCOTLAND:
Inverness-shire,
Rothiemurchus,
Watling
264C
(holo-
type
L.
vulpinum,
E).
—
SWEDEN:
Jamtland,
Borgsjo,
27
VIII
1993,
M.
E.
Noordeloos
9396',
Borgsjd
Julasen,
3
IX
1993,
M.
E.
Noordeloos
93156;
Borgsjo
Granbods&sen,
31
VIII
1993;
Borgsjo,
21
VIII
1999,
H.C.
den
Bakker
10;
Vasternorrland,
Ange,
25
VIII
1999,
G.
Redeuilh,
H.C.
den
Bakker
24.
Leccinum
vulpinum
can
readily
be
recognized
in
the
field
on
account
of
its
dark
red-
dish
brown
pileus
and
distinct
black
squamules
on
the
stipe.
The
intensity
of
the
blackish
discoloration
of
the
context
is
only
a
fraction
of
the
intensity
of
the
discoloration
as
seen
in
L.
versipelle
and
L.
aurantiacum,
which,
together
with
its
association
with
conifers,
can
help
to
facilitate
identification.
Leccinum
piceinum
25
is
another
species
that
has
been
described
from
coniferous
forests
in
Europe.
This
species
is
associated
with
Picea
and
was
originally
described
25)
Leccinum
piceinum
Pilat
&
Dermek,
Hribovite
huby
(1974) 153

540
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
vulpinum.
Fig.
8.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulo
cystidia.
Scale
bars
=
10
µm
resp.
1
cm.

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
541
from
central
Europe.
Most
collections
from
Sweden
that
were
collected
in
Picea
for-
est
and
therefore
identified
as
L.
piceinum
did
not
differ
in
their
pileus
colour
or
other
morphological
characters
from
L.
vulpinum.
The
collections
from
central
Europe
dif-
fered
from
L.
vulpinum
mainly
by
a
duller
pileus
colour,
a
character
that
is
also
used
by
Lannoy
&
Estades
(1995)
to
distinguish
L.
piceinum
from
L.
vulpinum.
The
holotype
of
L.
piceinum
was
not
sufficiently
well
preserved
to
make
a
morphological
study
possible,
however,
material
from
the
locus
typicus
did
not
differ
from
our
concept
of
L.
vulpinum,
except
for
the
pileus
colour.
In
an
Austrian
collection
of
L.
piceinum
we
amplified
an
ITS
sequence
type
that
differed
from
the
type
found
in
a
Norwegian
L.
vulpinum
(see
Den
Bakker
et
al.,
2004b).
Whether
the
central
European
collections
associated
with
Picea
with
an
aberrant
pileus
colour
represent
a
truly
genetically
distinct
taxon
remains
to
be
investigated.
Leccinum
vulpinum
is
probably
common
in
North
America.
A
Gapdh
sequence
of
a
collection
from
Manitoulin
Island
(Ontario,
Canada,
HC
den
Bakker
415
(L),
Genbank
accession
AY538793)
was
nearly
identical
to
the
Gapdh
sequence
of
L.
vulpinum
from
Norway.
Apparently
this
species
is
in
North
America
commonly
confused
with
L.
au-
rantiacum,
and
the
descriptions
of
Smith
&
Thiers
(1971)
of
L.
aurantiacum
and
plates
of
L.
aurantiacum,
L.
fibrillosum
26
and
L.
subtestaceum
27
in
Bessette
et
al.
(2000)
seem
to
actually
represent
L.
vulpinum
or
a
closely
related
species.
For
a
further
discussion
see
under
L.
aurantiacum.
4.
Leccinum
versipelle
(Fr.
&
Hök)
Snell
—
Fig.
9,
Plate
3
Boletus
versipellis
Fr.
&
Hok,
Bol.
Fung.
Gen.
111.
(1835)
13;
Leccinum
versipelle
(Fr.
&
Hok)
Snell,
Lloydia
7
(1944)
58.
—
Boletus
rufescens
Konrad,
Bull.
Mens.
Soc.
Linn.
Lyon
1
(1932)
151;
Leccinum
rufescens
(Konr.)
Sutara,
Ceska
Mykol.
43
(1989)
7.
—
Leccinum
testaceoscabrum
Singer,
Am.
Midi.
Naturalist
37
(1947) 123
[invalid,
no
Latin
diagnosis].
—
Boletus
percandidus
Vassilkov,
Sovetsk.
Bot.
2
(1944)
27
[invalid,
no
Latin
diagnosis];
Leccinum
percandidum
(Vassilkov)
Watling,
Trans.
Brit.
Mycol.
Soc.
43
(1960)691
[invalid,
basionym
not
validly
published];
Boletus
percandidus
Blum,
Bull.
Trimestriel
Soc.
Mycol.
France
85
('1969'
1970)
562;
Leccinum
percandidum
(Blum)
Lannoy
&
Estades,
Monogr.
Leccinum
Eur.
(1995)
112.
—
Leccinum
roseotinctum
Watling,
Notes
Roy.
Bot.
Gard.
Edinburgh
29
(1969)
267.
—
Leccinum
cerinum
M.
Korhonen,
Karstenia
35
(1995)
61.
—
Leccinum
callitrichum
Redeuilh,
Bull.
Trimestriel
Soc.
Mycol.
France
111
(1995)
169.
Neotype.
Sweden,
Jamtland,
Kali,
1
IX
2000,
H.C.
den
Bakker
070
(L,
designated
here).
Selected
icones.
Lannoy
&
Estades,
Mon.
Lecc.
Europe
(1995)
pi.
24,
pi.
31,
pi-
32,
pi.
33;
Estades
&
Lannoy,
Bull.
Trimestriel
Fdd.
Mycol.
Daupine-Savoie.
174
(3)
(2004)
74
(top).
Selected
description.
Watling,
Br.
Fung.
Fl.
1
(1970)
56.
Characteristics
—
A
slender
to
robust,
medium-sized
to
large
Leccinum;
pileus
light
brownish
to
orange,
sometimes
whitish,
light
brown
in
exsiccates.
Stipital
squamules
greyish
to
blackish,
also
in
young
fruit-bodies.
Mycorrhizal
with
Betula.
Pileus
80
-180
mm,
hemispherical
when
young,
convex
to
plano-convex
when
ma-
ture,
with
infiexed
margin
especially
in
young
fruit-bodies,
with
appendiculate
flaps,
exceeding
the
tubes
up
to
4
mm,
in
mature
fruit-bodies
marginal
flaps
often
absent
and
margin
entire,
yellowish
brown
(Mu.
10
YR
7/8),
orange
(2.5
YR
6/8)
to
brown
(5
YR
26)
Leccinum
fibrillosum
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
5
(1966)
165.
Fig.
14.
27)
Leccinum
subtestaceum
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
5
(1966)
145.
Figs.
5,6,7.

PERSOONIA
-
Vol.
18,
Part
4,
2005542
5/6),
sometimes
whitish
or
whitish
with
pinkish
or
apricot
tinges;
minutely
tomentose
to
fibrillose
squamulose.
Tubes
adnexed
to
emarginate,
segmentiform
to
ventricose,
8
-22
mm
long,
yellowish
white
to
brownish
grey,
violaceous
to
greyish
when
bruised.
Pores
c.
0.5
mm
in
diameter,
greyish
white
to
grey
ochre
(10
YR
7/3),
discolouring
brownish
when
bruised.
Stipe
70-200
x
10-45
mm,
cylindrical
to
clavate,
whitish,
greyish
white
or
yellowish
white,
sometimes
with
a
clear
blue
discoloration
in
the
stipe
base
when
handled,
lightly
to
densely
covered
with
fine
(sometimes
on
the
stipe
base
more
coarse)
greyish
to
blackish,
sometimes
whitish
squamules.
Context
white,
when
bruised
often
discolouring
greyish
or
blackish
with
a
violet
hue
in
the
pileus
and
upper
half
the
stipe,
often
discolouring
bluish
in
the
stipe
base.
Spores
(9.0—)
11.5-16.5
x
3.5-4.0(-5.0)
/mi,
Q
=
2.6-3.8,
Q
av
=
3.0-3.3;
fusiform
with
a
conical
apex,
with
a
distinct
suprahilar
depression,
pale,
purplish
brown
in
water.
Basidia
20-35
x
6.5-11.0
/mi,
clavate,
generally
4-spored.
Hymenocystidia
20-45
pm
x
7-11
/mi,
often
lageniform
and
hyaline
in
tubular
hymenium,
narrowly
utriform
or
clavate
(sometimes
with
an
obtuse
or
acuminate
apex),
hyaline
or
with
a
brownish
content
on
the
pore
surface.
Pileipellis
an
intricate
cutis
of
cylindrical
elements,
terminal
elements
obtuse,
40
-
90
x
5.0
-17.5
/mi,
pigment
granular
encrusted,
sometimes
form-
ing
small
globules,
pale
yellowish
brown.
Caulocystidia
(25-)40
-70
x
8.5
-15.0
/mi,
fusiform
to
utriform,
sometimes
conical
or
clavate,
largest
cystidia
generally
fusiform
or
utriform,
apex
usually
obtuse,
sometimes
acute,
pale
brownish,
with
greyish
content
in
KOH.
Clamp-connections
absent.
Habitat
&
distribution
—Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Betula
in
forests
and
heathland,
on
sandy,
slightly
acidic,
loamy
soils;
rather
rare
and
endangered
in
the
Netherlands
(Veerman,
2004),
more
common
in
Scandinavia
and
other
parts
of
Europe.
Collections
studied.
THE
NETHERLANDS:
Gelderland,
Leusderheide,
2001
,H.C.
den
Bakker
285;
Rhenen,
Buurtse
Berg,
8IX
1968,
W.
Wouters,
s.n.;
Winterswijk,
Nonnenven,
IX
2000,
R.
Chrispijn,
II.C.
den
Bakker
138.
—
FINLAND:
Inarilapland,
Utsjoki,
1995,
M.E.
Noordeloos
95702;
Enontekio
Lappi,
Kilpisjarvi,
1993,
M.
Korhonen
11452.
—
FRANCE:
Lozere,
Aumont-Aubrac,
Redeuilh,
2270P
(holotype
L.
callitrichum).
—
NORWAY:
Sogri
og
Fjordane,
Lasrdal,
12IX
2000,
OF64036.
—
SCOT-
LAND:
Inverness-shire,
Tomich,
Badger
Falls,
30
VIII
1957,
Watling
193c
(holotype
1
L.
roseotinctum
E).
—
SWEDEN:
Jamtland,
Borgsjo,
26
VIII
1999,
H.C.
den
Bakker
25;
Borgsjo,
IX
1999,
H.C.
den
Bakker
57;
Kali,
1
IX
2000,
H.C.
den
Bakker
70;
Kail,
1
IX
2000,
H.C.
den
Bakker
71;
Kali, 2
IX
2000,
H.C.
den
Bakker
74;
Vasternorrland,
delta
of
Indalsalven,
24
VIII
1999,
H.C.
den
Bakker
23;
Ostavall,
IX
1999,
H.C.
den
Bakker
4.
The
number
of
species
around
L.
versipelle
has
steadily
grown
in
the
past
decades.
Several
related
or
similar
species
have been
described,
such
as
L.
percandidum
with
a
white
or
whitish
pileus
that
becomes
pale
brownish
when
mature,
L.
roseotinctum,
with
an
initially
white
pileus
that
changes
to
pinkish
when
mature,
L.
cerinum
with
a
paler
orange
to
yellowish
pileus
and
pale,
yellowish
white
pores
in
young
basidiocarps,
and
L.
callit-richum
with
a
more
or
less
ochre
pileus
and
dark
brown
elements
in
the
pileipellis.
The
outcome
of
the
molecular
studies
(Den
Bakker
et
al.,
2004a,
2004b)
indicate,
however,
that
a
broad
morphological
species
concept
of
L.
versipelle
fits
better
with
these
results.
No
consistent
morphological
and
molecular
characters
sup-
port
the
recognition
of
species
like
L.
cerinum
and
L.
callitrichum.
The
collections
of
L.
roseotinctum
and
L.
percandidum
showed
sequences
that
were
(almost)
identical
to
the

543
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Fig.
9.
Leccinum
versipelle.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

544
PERSOONIA
-
Vol.
18,
Part
4,
2005
sequences
of
collections
of
normally
pigmented
L.
versipelle.
These
taxa
must
therefore
be
considered
as
less
pigmented
forms
of
L.
versipelle
without
formal
taxonomic
status.
Leccinum
atrostipitatum
A.H.
Sm.,
Thiers
&
Watling
is
an
originally
North
Ameri-
can
taxon
that
has
been
reported
repeatedly
from
Europe
(Engel,
1978)
based
on
the
observation
of
fruit-bodies
with
an
extremely
dark
and
very
dense
stipital
ornamenta-
tion.
Several
European
and
American
collections
identified
as
L.
atrostipitatum
have
been
studied,
but
no
significant
morphological
differences
between
these
collections
and
European
material
of
L.
versipelle
could
be
found.
ITS2
sequences
show
some
consistent
differences
between
North
American
and
European
collections.
We
therefore
think
that
these
differences
represent
infraspecific,
phylogeographic
differences
and
L.
atrostipitatum
and
L.
versipelle
are
actually
conspecific.
Further
North
American
collections
should
be
studied
and
sequenced
to
verify
this
assumption.
Given
the
fact
that
L.
versipelle
also
occurs
in
the
arctic
we
consider
L.
versipelle
a
Betula-
associate
with
a
circumboreal
distribution.
SUBSECTION
FUMOSA
A.H.
SM.,
Thiers
&
Watling, Michigan
Bot.
6
(1967)
138
Pileus
margin
entire;
context
white
turning
black when
bruised,
rarely
unchanging.
Widespread
throughout
the
Northern
Hemisphere.
Type
species:
Leccinum
olivaceoglutinosum
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
6(1967)
140.
5.
Leccinum
duriusculum
(S.
Schulz.)
Singer
—
Fig.
10,
Plate
4
Boletus
duriusculus
S.
Schulz.
in
Fr.,
Hymenomyc.
Eur.
(1874)
515;
Leccinum
duriusculum
(S.
Schulz.)
Singer,
Amer.
Midi.
Naturalist
37
(1947)
122.
—
Leccinum
duriusculum
f.
robustum
Lannoy
&
Estades
in
Doc.
Mycol.
24
(94)
(1994)
18.
—
Leccinum
nigellum
Redeuilh,
Bull.
Trimestriel
Soc.
Mycol.
France
111(3)
(1995)
174.
Holotype:
not
existent;
type
locality:
Slavonia
(Croatia).
Selected
icones.
Bon,
Mushr.
Toadst.
(1987)
41;
Breitenb.
&
Kranzl.,
Pilze
Schweiz
3
(1991)
70,
pi.
32;
Engel,
Rauhstielrohrlinge
(1978)
pi.
5a;
Munoz,
Fungi
non
del.
13
(2000)
pi.
11,
12
(as
f.
robustum
);
R.
Phillips,
Paddest.
Schimm.
(1981)
212;
Pilat
&
Dermek,
Hrfvobite
Huby
(1974)
pi.
75.
Selected
descriptions
&
illustrations.
Breitenb.
&
Kranzl.,
Pilze
Schweiz
3
(1991)
70,
pi.
32;
Engel,
Rauhstielrohrlinge
(1978)
33, 34;
Estades
&
Lannoy,
Bull.
Trimestriel
Fed.
Mycol.
Daupine-Savoie
174
(2004)
70
(as
f.
robustum
);
Munoz,
Fungi
non
del.
13
(2000)
30-32;
Watling,
Br.
Fungus
Fl.
I
(1970)
50.
Characteristics
—
Medium
to
large-sized
and
sturdy
Leccinum
with
dark
grey-brown
to
blackish
pileus
with
an
appendiculate
margin
without
marginal
flaps;
pores
whitish
to
pale
ochre;
stipe
white
with
dark
brown
to
blackish
squamules,
which
sometimes
ag-
glutinate
and
form
a
net-like
structure
in
upper
part;
context
white,
turning
black when
bruised,
rarely
unchanging;
pileipellis
cutis-like.
Associated
with
Populus
tremula,
P.
alba
and
various
planted
Populus
hybrids.
Pileus
40-150(-220)
mm,
hemispherical,
then
convex,
finally
sometimes
with
slightly
flattened
centre,
very
variable
in
colour
from
rather
pale
grey-brown
with
slight
violaceous
tinge
to
rather
dark
greyish
or
reddish
brown
(Mu.
10
YR
4-7/4;
7.5
YR
7/4-4/2),
minutely
fibrillose-felted
to
very
minutely
appressedly squamulose
all
over,

545
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
often
very
finely
cracked
with
age,
forming
very
small
fibrillose-felted
patches,
initially
concolorous
with
context,
but
later
contrasting
with
paler
context.
Tubes
adnexed
to
almost
free,
broadly
ventricose,
up
to
25
mm
long,
creamy
white
then
greyish
buff;
pores
rounded,
about
0.3-0.5
mm
in
diameter,
brownish
white
(5
Y
7-6/3),
turning
brown
when
bruised.
Stipe
80-170
x
14-20
mm,
cylindrical
with
slightly
tapering
apex
and
rounded
to
subclavate
base,
solid,
entirely
minutely
squamulose
with
fine
greyish
to
blackish
squamules,
which
become
larger
towards
base,
often
arranged
in
longitudinal
rows
or
ridges;
sometimes,
especially
in
upper
part
of
stipe,
merging
into
a
reticulate
pattern,
strongly
contrasting
with
pallid,
almost
white
stipital
surface.
Context
white
in
pileus
and
upper
part
of
stipe,
frequently
tinged
yellow-green
in
lower
part
of
stipe,
slowly
turning
violaceous
pink
when
cut,
then
spot-wise
turning
darker
greyish-vio-
laceous
black;
in
base
of
stipe
locally
turning
yellow-green
to
blue-green.
Spore
print
olivaceous-yellow
brown
(2.5
Y 6/6
tending
to
10
YR
5/6).
Spores
(11.5
—)12.0
—15.5
x
4.5-6.0
//m,
on
average
13.0-14.0
x
4.8-5.0
//m,
Q=2.3-3.3,
Q
av
=2.4-2.7;
fusiform with
conical
apex,
with
distinct
suprahilar
de-
pression,
relatively
thick-walled,
pale
brown
in
water.
Basidia
20-35
x
6.0-9.0
pm,
clavate,
4-spored.
Hymenial
cystidia
abundant
on
edge
and
sides
of
the
pores,
20-75
x
5.5-17
x
2.0-4.0
pm,
lageniform,
often
with
rather
long,
tapering
neck,
colourless
or
with
brown,
intracellular-granular
pigment.
Pileipellis
cutis-like,
composed
of
septate
hyphae,
terminal
elements
12-90(-120)
x
2.5-7.5
pm,
irregularly
cylindrical
with
intracellular,
granular
pigment.
Stipitipellis
a
cutis
of
narrow,
cylindrical,
3.0-9.0
pm
wide
hyphae,
with
small
to
fairly
large
caulohymenial
clusters
of
basidia
and
cystidia,
forming
the
squamules
on
the
stipital
surface.
Caulocystidia
25
-110
x
5.0-12
x
2.0-6.5
//m,
lageniform
or
fusiform,
thin-
or
thick-walled,
colourless
or
with
brown
granular
intracellular
pigment.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Populus,
especially
with
P.
alba
and
P.
tremula,
but
also
with
various
Populus
cultivars,
often
along
roadsides,
preferably
on
calcareous
clay,
sometimes
on
sand
or
loam;
in
the
Netherlands
locally
rather
common,
elsewhere
rare,
mainly
inN
and
F,
very
rare
in
L,
E,
Z,
D
and
K.
Rare,
but
widespread
in
Europe.
Collections
studied.
THE
NETHERLANDS:
Utrecht,
Werkhoven,
IX
2004,
J.
Wisman
s.n.;
Gelderland,
Beesd,
10
VIII
1973,
C.
Bas
5865;
Neerijnen,
Kasteelbos,
16
VIII
1970,
F.
Tjallingii
s.n.;
Lingebos,
17
VII
1993,
C.
Bas
8876;
Zuid-Holland,
Wassenaar,
8
VII
2004,
H.C.
den
Bakker
s.n.
—
FRANCE:
Foret
de
Vibraye,
11
IX
1979,
Redeuilli
4676
P
(holotype
L.
nigellum.
,PC).
Leccinum
duriusculum
f.
robustum,
,
which
according
to
Lannoy
&
Estades
(1995)
differs
by
more
robust
basidiocarps,
is
not
recognized
here.
A
continuum
in
size
and
shape
between
f.
duriusculum
and
f.
robustum
has
been
observed.
The
last-mentioned
form,
however,
seems
to
prefer
the
hybrid
Populus
x
cinerea,
whereas
the
more
slender
form
often
grows
with
Populus
tremula.
Exceptions
on
this
pattern
of
host
association
have been
found,
and
hence the
forms
are
not
given
a
formal
status
here.
Leccinum
nigellum
differs
from
L.
duriusculum
mainly
in
the
dark
(almost)
blackish
colour
of
the
pileus.
Molecular
data
(Den
Bakker
et
al.,
2004b)
show
that
it
is
closely
related
to
or
conspecific
with
L.
duriusculum.
Given
the
species
concept
applied
in
this
study
(a
species
must
differ
from
a
related
species
in
at
least
two
independent
characters
and
must
be
monophyletic),
we
consider
L.
nigellum
a
dark
form
of
L.
duriusculum.

PERSOONIA
-
Vol.
18,
Part
4,
2005
546
Leccinum
duriusculum.
Fig.
10.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

547
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
SUBSECTION
SCABRA
Pilát
&
Dermek,
Hribovite
Huby
(1974)
137
Pileal
margin
entire.
Context
whitish,
showing
bluish,
pinkish
or
reddish
discolora-
tions
when
bruised,
or
no
discoloration
at
all,
sometimes
light
greyish
after
several
hours.
Mycorrhizal
with
Betula.
Throughout
the
Northern
Hemisphere,
following
the
distribution
of
Betula.
Type
species:
Leccinum
scabrum
(Bull.:
Fr.)
S.F.
Gray
6.
Leccinum
scabrum
(Bull.:
Fr.)
Gray
—
Fig.
11,
Plate
5
Boletus
scaber
Bull.,
Herb.
Fr.
(1782)
pi.
132;
Boletus
scaber
Bull.
:
Fr.,
Syst.
mycol.
1
(1821)
393;
Leccinum
scabrum
(Bull.:
Fr.)
Gray,
Nat.
Arr.
Brit.
PI.
1
(1821)
647.
—
Leccinum
roseofractum
Watling,
Notes
Roy.
Bot.
Gard.
Edinburgh
28
(1968)
313.
—
Boletus
avellaneus
Blum,
Bull.
Trimes-
triel
Soc.
Mycol.
France
85
('
1969'
1970)
560;
Leccinum
avellaneum
(Blum)
Bon. Doc.
Mycol.
9(35)
(1979)
41.
—
Leccinum
rigidipes
P.D.
Orton,
Trans.
Brit.
Mycol.
Soc.
91
(1988)
560.
—
Krombholzia
scabra
f.
cinnamomea
Valsilkov,
Trudy
Bot.
Inst.
2
(10)
(1956)
370,
non
Leccinum
cinnamomeum
A.H.
Sm.,
Thiers
&
Watling
1967;
Leccinum
subcinnamomeum
Pilat
&
Dermek,
Ziva
17
(2)
(1969)
58.
Iconotype.
Bull.,
Herb.
France:
pi.
132.
1782.
Epitype.
J.
van
Brummelen
1+2,
along
N12,
vicinity
of
Versailles,
Guyancourt,
France
(L,
desig-
nated
here).
Selected
icones.
Lannoy
&
Estades,
Monogr.
Leccinum
(1995)
pi.
4;
Marchand,
Champ.
Nord
Midi
2
(1973)
pi.
169;
R.
Phillips,
Paddest.
Schimm.
(1993)
213;
Ryman
&
Holmasen,
Pilze
(1992)
231;
Lannoy
&
Estades,
Monogr.
Leccinum
(1995)
pi.
16
(as
L.
rigidipes)
;
R.
Phillips,
Paddest.
Schimm.
(1993)
213
(als
L.
oxydabile).
Selected
descriptions.
Lannoy
&
Estades,
Monogr.
Leccinum
(1995)
62-64;
J.
Schreiner,
Beih.
Z.
Mykol.
9(1999)
27-32.
Characteristics
—
Habit
slender
to
robust,
small
to
medium-sized;
pileus
ranging
from
light
brownish
to
dark
brown;
stipital
squamules
light
brownish
to
blackish,
coarse
in
the
lower
half
of
the
stipe
and
usually
fine,
almost
glandular
in
the
upper
half
of
the
stipe;
context
always
without
bluish
discolorations.
Mycorrhizal
with
Betula
in
(dry)
acidic
habitats.
Pileus
52-130
mm,
convex
at
first,
expanding
to
plano-convex,
with
entire,
deflexed
or
inflexed
margin,
not
or
only
very
slightly
(<
1mm)
extending
over
tubes,
yellowish
brown
to
dark
brown
(Mu.
10
YR
6/6,
7.5
YR
3-6/4);
minutely
tomentose,
breaking
up
into
minute,
appressed
squamules
with
age,
somewhat
viscid
when
moist.
Tubes
broadly
adnexed,
8-19
mm
long,
segmentiform
to
subventricose
when
mature,
whitish
when
young,
brownish
grey
when
mature.
Pores
greyish
white,
often
with
brownish
spots,
discolouring
brownish
when
bruised.
Stipe
82-145
x
12-55
mm,
cylindrical
to
clavate,
whitish,
often
more
brownish
towards
apex,
entirely
covered
with
blackish
to
greyish,
sometimes
pale
brownish
squamules,
fine
at
apex,
becoming
gradually
coarser
towards
base,
sometimes
agglutinating
and
almost
forming
a
network.
Context
whitish,
not
changing
colour
when
bruised
or
at
most
discolouring
pinkish
or
reddish
(K.
&
W.
9A2),
finally
often
brownish
or
slightly
greyish
after
several
hours.
Spores
(13.5-)
14.5-19.0(-22.0)x
(4.0-)5.0-6.5
//m,
Q
=
2.5-3.5,
Q
av
=
(2.9-)3.0-3.3,
fusiform
with
a
suprahilar
depression.
Basidia
30-40
x
10.5
-12.5
/mi,
4-spored
(sometimes
3-spored).
Hymenocystidia
30-50
x
5.5-7.5
m,
fusiform
to

548
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
scabrum.
Fig.
11.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

549
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
lageniform,
cylindrical
to
clavate
or
lageniform,
apexobtuse
or
acuminate.
Caulocystidia
(35-)
40-70
(-
90)
x
9.0-18.5
pm,
lageniform
to
clavate.
hyaline
or
with
a
greyish
brown
content
when
mounted
in
KOH.
Pileipellis
a
dense
to
loosely
intricate
cutis,
composed
of
slender,
hyaline
to
brownish
intracellularly
pigmented,
or
granular
incrusted
hyphal-
elements,
3.5
—7.0(—
8.0)
pm
in
diameter,
mixed
with
broadly
elongated
hyphal
elements,
(7.5-)8.0-14.0(-16.0)
pm
in
diameter,
sometimes
also
with
broad,
clearly
articulated,
hyaline
hyphal
elements,
(8.0-)9.0-13.5(-17.0)
pm
in
diameter,
terminal
elements
often
conical.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal
with
Betula,
often
found
in
lawns
or
oligotrophous
Festuca
rubra
/Deschampsia
flexuosa
vegetations
on
slightly
acidic,
dry
sandy
soils;
common
in
the
Netherlands,
widespread
and
common
in
Europe.
Probably
a
species
with
a
circumboreal
distribution.
Collections
studied.
THE
NETHERLANDS:
Drenthe,
Emmen,
2
X
1999,
H.C.
den
Bakker
44;
Nieuw
Amsterdam,
4
X
1999,
H.C.
den
Bakker
50;
Borger,
vicinity
of
cottage
park
'Land
van
Bartje',
5
X
1999,
H.C.
den
Bakker
54;
Gieten,
Boekweitveentje,
2001,
R.
Enzlin
01-009;
Hoogeveen,
3
X
1999,
Leg.
B.
de
Vries,
H.C.
den
Bakker
48;
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
Eiland
van
Rolvers,
23
X
1999,
H.C.
den
Bakker
62;
Zuid-Holland,
Wassenaar,
Meijendel,
17
X
1999,
H.C.
den
Bakker
58;
Voorschoten,
30
IX
1999,
C.
Bas,
H.C.
den
Bakker
42;
Utrecht,
Zeist,
Soesterberg,
6
X
2000,
H.C.
den
Bakker
124.
—
FRANCE:
Versaille,
27
IX
2001,
Van
Brummelen
(epitype.
L.
scabrum,
L);Orne,
Foret
de
Belleme,
281X2001,
Van
Brummelen.
—
NORWAY:
Sognog
Fjordane,
Sogndalsfjpra,
8IX
2000,
H.C.
den
Bakker
95;
Sogn
og
Fjordane,
Sogndalsfjpra,
8IX
2000,
H.C.
den
Bakker
91;
Sogn
og
Fjordane,
Sogndalsfjpra,
IX
2000,
H.C.
den
Bakker
87.
—
SCOTLAND:
Inverness-shire,
Aberarder
Forest,
13IX
2001.
H.C.
den
Bakker
239;
Insh
Marshes,
12IX
2001,
H.C.
den
Bakker
231;
Insh
Marshes,
12IX
2001
,H.C.
den
Bakker
237;
Abernethy
Forest,
31
VIII
1972,
P.
Orton
1
4399
(holotype
L.
rigidipes,
E);
Ross
&
Cromarty,
Loch
Maree,
R.
Watling
4944
(holotype
L.
roseofractum.
E).
—
SWEDEN:
Jamtland,
Borgsjo,
24
VIII
1999,
H.C.
den
Bakker
22;
Are,
4IX
2000,
H.C.
den
Bakker
80.
—
WALES:
South
Glamorgan,
Merthyr-mawr
sand
dunes,
A.
Hills
2003076.
Leccinum
scabrum
can
usually
be
distinguished
from
other
species
of
subsection
Scabra
by
a
combination
of
the
following
characters:
-
the
stipital
ornamentation
is
usually
coarse
in
the
lower
part
of
the
stipe
and
with
fine,
sometimes
almost
'glandular'
squamules
in
the
upper
half
of
the
stipe.
-
bluish
discolorations
of
the
context
are
always
absent.
-
the
presence
of
remarkably
large
(up
to
92
pm
long),
usually
clavate
caulocystidia.
-
the
pileipellis
is
a
loose
cutis
composed
of
a
mixture
of
slender
and
broad,
sometimes
elliptical,
elements.
Leccinum
scabrum
generally
occurs
in
drier
habitats
than
other
species
of
subsec-
tion
Scabra.
Thanks
to
monographs
like
the
one
of
Lannoy
&
Estades
(1995)
there
is
a
general
consensus
about
the
identity
of
L.
scabrum.
The
current
concept,
however,
of
L.
scabrum
is
wider
than
that
of
Lannoy
&
Estades,
and
includes
also
L.
roseofractum,
L.
avellaneum
and
L.
rigidipes,
thus
including
also
forms
with
discolouring
context.
As
such,
L.
scabrum
is
a
fairly
variable
species,
and
L.
avellaneum
and
L.
roseofractum
may
be
considered
the
extremes
in
a
series
of
lighter
to
darker
forms.
Gradual
differences
were
also
found
in
the
discoloration
of
the
context
when
bruised,
ranging
from
not
re-
acting
to
discolouring
pinkish
to
almost
reddish.
To
consolidate
the
presently
accepted
interpretation
of
L.
scabrum
an
epitype
accompanying
the
iconotype,
originating
from
the
neighbourhood
of
Paris,
is
designated
here.

550
PERSOONIA
-
Vol.
18,
Part
4,
2005
In
Europe,
the
slender
spores
(Q
av
a
3.0)
can
help
to
separate
L.
scabrum
from
L.
rotundifoliae.
However,
the
average
Q-value
of
collections
identified
as
L.
scabrum
from
eastern
North
America
is
lower
than
the
average
Q-value
in
Europe,
viz.
2.8
in-
stead
of
3.0.
Therefore
the
average
Q-value
can
probably
not
be used
in
North
America
to
discriminate
this
species
from
L.
rotundifoliae.
In
North
America
the
large
caulo-
cystidia
and
the
irregular
pileipellis
structure
can
be used
to
distinguish
L.
scabrum
from
L.
rotundifoliae.
According
to
Watling
(1968)
L.
roseofractum
differs
from
L.
scabrum
by
the
more
robust
habit,
dark-coloured
pileus,
the
dense
black
stipital
ornamentation
and
flesh
changing
distinctly
red.
Collections
were
made
of
L.
roseofractum
in
Scotland,
Norway
and
Canada.
The
European
collections
matched
all
of
Watling's
criteria
for
L.
roseo-
fractum;
the
Canadian
collection,
however,
had
a
slender
habit,
but
a
strong
reddish
discoloration,
an
almost
blackish
pileus
and
dark
blackish
stipital
ornamentation.
The
three
collections
did
not
form
a
separate
clade
in
our
molecular
phylogenetic
analysis
(Fig.
2),
but
were
found
dispersed
between
accessions
of
L.
scabrum.
Microscopically
these
collections
and
the
type
collection
of
L.
roseofractum
were
not
different
either
from
collections
identified
as
L.
scabrum.
Therefore
L.
roseofractum
is
considered
a
synonym
of
L.
scabrum.
The
type
material
and
original
description
of
Orton
(1988)
of
L.
rigidipes
is
remi-
niscent
of
L.
scabrum
in
almost
every
morphological
character,
except
for
the
presence
of
short
hyphal
elements
(interpreted
as
cylindrocysts
by
Lannoy
&
Estades,
1995)
in
the
pileipellis.
Specimens
with
abundant
cylindrocysts,
identified
as
L.
rigidipes,
were
molecularly
identical
with
collections
without
cylindrocysts.
Moreover,
we
noted
that
cylindrocysts,
though
in
low
numbers,
are
often
present
in
the
pileipellis
of
L.
scabrum.
In
conclusion,
L.
rigidipes
is
also
considered
synonymous
with
L.
scabrum.
The
name
L.
subcinnamomeum
is
validly
published,
since
Pilat
and
Dermek
referred
to
Krombholzia
scabra
f.
cinnamomea
Vasilkov
as
being
the
basionym
of
this
taxon.
Kromholzia
scabra
f.
cinnamomea
Vasilkov
is
validly
published
and
is
not
the
basionym
of L.
cinnamomeum
A.H.
Sm.,
Thiers
&
Watling
(1967).
No
type
material
of.
L.
subcin-
namomeum
seems
to
be
designated.
Material
of
this
species
in
PRM
fits
perfectly
in
our
morphological
concept
of
L.
scabrum.
Most
descriptions
and
plates
in
the
literature
(Engel,
1978;
Dahnke,
1993;
Bresinsky,
1996)
also
refer
to
a
taxon
close
to
L.
scabrum
with
a
reddish
brown
pileus.
7.
Leccinum
melaneum
(Smotl.)
Pilát
&
Dermek
—
Fig.
12,
Plate
6
Boletus
scaber
var.
melaneus
Smotl.,
Mykol.
Shorn.
16
(1951)
17;
Leccinum
melaneum
(Smotl.)
Pildt
&
Dermek,
Hrfbovite
huby
(1974)
145;
Leccinum
scabrum
var.
melaneum
(Smotl.)
Dermek,
Fung.
rar.
Ic.
col.
16
(1987)
17.
Selected
icones.
Dahncke,
1200
Pilze
(1993)
pi.
98.
Characteristics
—
A
dark,
medium-sized
to
large
Leccinum.
Pileus
brown
to
blackish
brown;
stipe
greyish
to
blackish
with
dense
blackish
stipital
squamules,
stipe
usually
strongly
clavate;
context
without
any
bluish
discolorations.
Mycorrhizal
with
Betula.
Pileus
50-130
mm,
convex
at
first,
expanding
to
plano-convex
with
age,
often
irregularly
shaped,
brown
to
dark
brown
(Mu.
7.5
YR
3/4),
sometimes
with
lighter
coloured
spots;
very
finely
tomentose,
becoming
subviscid
with
age.
Tubes
narrowly

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
551
Leccinum
melaneum.
Fig.
12.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

PHRSOONIA
-
Vol.
18,
Part
4,
2005552
to
broadly
adnexed,
8-19
mm
long,
whitish
when
young,
greyish
to
brownish
at
ma-
turity.
Pores
greyish
white,
often
with
brownish
spots,
brownish
when
bruised.
Stipe
90-145
x
21-55
mm,
cylindrical
to
(broadly)
clavate,
greyish
to
blackish,
entirely
covered
with
squamules;
squamules
fine
near
the
apex
of
the
stipe,
gradually
becoming
coarser
towards
base,
in
basal
part
often
agglutinated
and
forming
a
kind
of
reticulum.
Context
white,
when
bruised
often
discolouring
pinkish
(K.
&W.
9A2),
often
brownish
after
several
hours.
Spores
14.0-19.0
x
5.0-6.0
//m,
Q
=
2.5-3.5,
Q
av
=
2.9-3.2,
fusiform
with
a
su-
prahilar
depression.
Basidia
20-30
x
11.5-13.0
//m,
4-spored,
sometimes
2-spored.
Hymenocystidia
25-50
x
7.5-11.5
pm,
lageniform,
clavate
with
a
mucronate
apex
or
utriform.
Pileipellis
a
loose
intricate
cutis
of
cylindrical,
sometimes
bifurcate,
elements
9.0-11,5(-15.0)
pin
i
wide,
with
dark
brown
intracellular
pigment.
Caulocystidia
35-60
x
12.5-18.5
pm,
fusiform,
utriform
or
clavate.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Betula
on
sandy
soils.
Known
from
four
localities
in
the
Netherlands
(Boekweitveentje,
Gieten;
Boswachterij
Gees;
Amsterdamse
Waterleiding
Duinen,
Wisselsche
Veen,
Epe),
susceptible
(Veerman
2004).
Distribution
in
Europe
unclear,
because
of
confusion
with
dark
forms
of
L.
scabrum.
Collections
studied.
THE
NETHERLANDS:
Drenthe,
Gieten,
Boekweitveentje,
3
X
1999,
H.C.
den
Bakker
47;
Boswachterij
Gees,
5
X
1999,
H.C.
den
Bakker
53;
Gelderland,
Epe,
25
X
2003,
Boom-
sluiter
s.n.;
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
Eiland
van
Rolvers,
2IX
1999,
H.C.
den
Bakker
33.
Leccinum
melaneum
does
not
differ
morphologically
much
from
L.
scabrum,
except
for
the
fact that
the
stipital
surface
generally
is
greyish
or
blackish
in
this
species,
as
opposed
to
whitish
to
brownish
in
L.
scabrum.
Three
of
the
four
collections
also
show
an
unusually
broad
stipe
base
(giving
the
stipe
a
broadly
clavate
shape)
and
a
slightly
malformed
pileus.
Molecular
data
indicate
L.
melaneum
might
be
a
hybridogenic
taxon
(Den
Bakker
et
al„
in
prep).
In
PRM
several
collections
identified
as
L.
melaneum
by
Pilat
are
present,
however,
as
far
as
we
know
no
type
material
for
L.
melaneum
has
been
designated.
Thismay
mean
L.
melaneum
has
never
been
validly
published.
We
therefore
see
L.
melaneum
as a
'preliminary'
name,
until
the
taxonomic
and
nomenclatural
status
of
this
taxon
has
been
clarified.
8.
Leccinum
rotundifoliae
(Singer)
A.H.
Sm.,
Thiers
&
Watling
—
Fig.
13,
Plate
7
Krombholzia
rotundifoliae
Singer,
Schweiz.
Z.
Pilzk.
16(1938)
148;
Boletus
rotundifoliae
(Singer)
S.
Lundell,
Fungi
Exs.
Suec.,
praesert.
Upsal.
(1959)
2615;
Leccinum
rotundifoliae
(Singer)
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
6
(1967)
128.
Leccinum
pulchrum
Lannoy
&
Estades,
Doc.
Mycol.
21
(82)
(1991)
3.—
L.
scabrum
subsp.
tundrae
Kallio,
Rep.
Kevo
Subarct.
Res.
Stat.
12
(1975)
25-27.
—
Leccinum
pulchrum
f.
fuscodiscum
Lannoy
&
Estades,
Doc.
Mycol.
21(82)(1991)
5.
Excluded.
L.
rotundifoliae
sensu
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
6
(1967)
128;
sensu
Kallio,
Rep.
Kevo
Subarct.
Res.
Stat.
12
(1975)
25-27
(=
L.
holopus).
Neotype
(selected
by
Lannoy
&
Estades,
1995).
Fungi
exsiccati
Suecici
2615,
Sweden,
Norrbotten,
Abisko,
27
VII
1943,
Nannfeldt
6771
(PC).
Selected
icones.
Lannoy
&
Estades,
Mon.
Leccinum
(1995)
pi.
2
(as
L.
pulchrum)
),
pi.
3
(as
L.
pulchrum
f.
fuscodiscum),
pi.
7;
Estades
and
Lannoy,
Bull.
Trimestriel
Fed.
Mycol.
Dauphine-
Savoie
174
(2004)
62
(as
L.
pulchrum).

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
553
Fig.
13.
Leccinum
rotundifoliae.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

554
PERSOONIA
-
Vol.
18,
Part
4,
2005
Selected
descriptions.
Kallio,
Rep.
Kevo
Subarct.
Res.
Stat.
12
(1975)
25-27
(as.
L.
scabrum
subsp.
tundrae)
Characteristics
—
A
slender
to
robust,
small
Leccinum.
Pileus
pale
brownish
or
dark
brown
with
lighter
regions;
stipital
squamules
brownish
to
blackish,
often
warty,
almost
flocculose
in
the
lower
half
of
the
stipe;
context
always
without
bluish
discolorations.
Mycorrhizal
with
Betula
in
boreal/subalpine
to
arctic/alpine
habitats.
Pileus
25-90
mm,
hemispherical
to
convex,
expanding
with
age,
with
inflexed,
entire
margin,
not
exceeding
the
tubes
or
at
most
up
to
1
mm,
pale
yellowish
brown,
light
brown
(Mu.
10
YR
5/2)
to
very dark
brown
or
grey
brown
(10
YR
6/4),
then
often
mottled;
surface
fibrillose-tomentose,
sometimes
appearing
velutinous,
frequently
ir-
regularly
cracked;
slightly
viscid
when
old.
Tubes
adnexed
to
narrowly
adnate,
8-21
mm
long,
broadly
convex
to
ventricose,
creamy
white
to
greyish
white.
Pores
yellowish
white
to
brownish
white,
discolouring
brownish
when
bruised.
Stipe
40-100
x
7-25
mm,
subclavate
to
clavate,
whitish;
entirely
covered
with
squamules,
squamules
either
whitish
and
changing
to
brownish
with
maturity
or
blackish,
fine
near
the
apex,
gradually
becoming
more
coarse/flocculose
towards
stipe
base.
Context
whitish,
when
bruised
unchanging
or
discolouring
pinkish,
especially
in
apex
of
stipe.
Spores
(11.5-)
13.5-17.5
(-20.0)
x
4.5
-7.0
pm,
Q=2.2-3.2,
Q
av
=
2.5-2.7(-2.9).
Basidia
23.5-33.0
x
11.0-12.5
pm,
clavate.
Pileipellis
a
rather
regular
intricate,
cutis;
elements
30-95
x
3.0-8.5
/on,
narrowly
cylindrical,
hyaline
and
granular
incrusted
or
brownish
and
then
often
also
granular
incrusted.
Caulocystidia
35-60
x
11.0-20.5,
utriform,
lageniform
or
fusiform,
content
hyaline
or
pale
greyish
brown
when
mounted
in
KOH.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Betula
in
wet
to
dry
habitats
(ranging
from
tundra
to
wet
birch
forest);
not
found
in
the
Netherlands,widespread
in
boreal,
arctic
and
alpine
regions,
probably
following
a
cir-
cumpolar
distribution.
Exact
distribution
in
subalpine
regions
at
lower
latitudes
unclear
because
of
insufficiently
known
morphological
variability
(Den
Bakker
et
al.
in
prep.).
Collections
studied.
CANADA:
Labrador,
Schefferville,
4IX
1984,
Malloch
4-9-84/1.
—
FINLAND:
Enontekion
Lappi,
Kilpisjarvi,
M.
Korhonen
11429.
—
FRANCE:
Isere,
Massif
de
Belledonne,
Prabert,
24X1989,
Estades
891024/3
(holotype
L.
pulchrum,
LIP);
Massif
de
Belledonne,
Les
Seiglieres,
28
IX
1989,
Estades
89.09.284
bisA
(holotype
L.
pulchrum
F
.fuscodiscum,
LIP).
—
GREENLAND:
Kel-
ly
ville,
14
VIII
2000,
L.
Jalink
&
M.
Nauta
8263;
Sisimiut,
VIII
2000,
L.
Jalink
&
M.
Nauta
8254;
Sisimiut,
19
VIII2000,
L.
Jalink
&
M.
Nauta
8399;
Sisimiut,
VIII2000,
L.
Jalink
&
M.
Nauta
8359.
—
NORWAY:
Jotunheimen,
6IX
2000,
H.C.
den
Bakker
86.
—
SWEDEN:
Jamtland,
Borgsjo,
23
VIII
1999
,H.C.
den
Bakker
16.
Leccinum
rotundifoliae
is
a
species
of
arctic,
alpine,
boreal
and
subalpine
habitats.
Pale-coloured
fruit-bodies
can
be
distinguished
from
L.
holopus
by
the
absence
of
blu-
ish
discolorations
in
the
context
of
the
stipe
base,
and
the
absence
of
greenish
bluish
tinges
in
the
pileus
of
older
fruit-bodies.
In
addition
to
differences
in
colour
there
seems
to
be
an
ecological
difference
in
arctic
regions.
Leccinum
rotundifoliae
is
a
species
of
dry
habitats,
whereas
L.
holopus
prefers
wet
habitats.
In
subalpine
and
boreal
regions
L.
rotundifoliae
can
also
be
found
in
wetter
habitats.
The
concept
of
L.
rotundifoliae
of
Smith
et
al.
(1967)
differs
from
the
original
de-
scription
of
L.
rotundifoliae.
Singer's
original
diagnosis
states
that
the
context
does
not

555
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
change
colour
when
bruised.
Smith
et
al.
(I.e.)
described
a
pale,
small
species
with
a
bluish
discoloration
of
the
context
in
the
stipe
base.
Collections
from
arctic
environ-
ments
fitting
the
concept
of
Smith
et
al.,
thus
with
a
bluish
discoloration
in
the
stipe
base,
appeared,
however,
in
phylogenetic
analyses
in
clades
together
with
L.
holopus
(see
Figs.
2,3),
and
not
in
the
L.
rotundifoliae
clade.
In
fact,
the
concept
of
Smith
et
al.
actually
represents
an
arctic
phenotype
of
L.
holopus.
These
two
different
concepts
of
L.
rotundifoliae
have
coexisted
for
some
time
in
the
literature
and
have
generated
abundant
confusion.
Kallio
(1975),
who
was
among
the
first
to
realise
that
two
different
arctic
species
were
involved,
followed
the
interpretation
of
L.
rotundifoliae
by
Smith
et
al.
when
describing
L.
scabrum
subsp.
tundrae.
He
listed
a
number
of
morphological
and
ecological
differences
between
L.
rotundifoliae
and
L.
scabrum
subsp.
tundrae,
from
which
is
apparent
that
his
concept
of
L.
rotundifoliae
fits
that
of
Smith
et
al.,
while
the
description
of
L.
scabrum
subsp.
tundrae
perfectly
fits
the
present
concept
of
L.
rotundifoliae.
Leccinum
rotundifoliae
is
generally
considered
a
pale
brown
to
almost
whitish
spe-
cies.
However,
specimens
with
a
dark
pileus
and
dark
stipital
squamules
were
also
sequenced.
The
first
of
these
specimens
to
be
sequenced
(a
collection
from
Borgsjo,
Jamtland,
Sweden)
was
erroneously
called
L.
scabrum
(Den
Bakker
et
al.,
2004a)
based
on
the
minor
sequence
divergence
in
ITS
between
L.
scabrum
and
L.
rotundifoliae.
Analysis
of
an
additional
gene
(
Gapdh
),
however,
supported
the
recognition
of
L.
rotun-
difoliae
as a
separate
species.
When
type
material
of
L.
pulchrum
was
sequenced,
these
accessions
surprisingly
appeared
in
the
L.
rotundifoliae-clade.
Upon
closer
inspection
of
the
morphology
and
a
comparison
of
photographs
of
dark
specimens
of
L.
rotun-
difoliae
from
Greenland
and
Sweden
with
the
picture
of
L.
pulchrum
in
Estades
&
Lannoy
(2004:62)
these
pictures
show
a
perfect
resemblance.
Leccinum
pulchrum
is
therefore
reduced
into
the
synonymy
of
L.
rotundifoliae.
This
demonstrates
that
L.
rotundi-
foliae
not
only
occurs
in
Northern
Europe,
but
also
in
the
Alps.
One
accession
from
the
French
lowlands
which
was
identified
as
L.
pulchrum,
appeared,
however,
in
the
L.
scabrum
clade.
Morphological
data
confirmed
that
the
collection
belongs
to
L.
scabrum,
and
the
name
that
was
initially
provided
(
L.
pulchrum)
must
therefore
be
considered
a
misapplication.
In
their
description
of
L.
pulchrum,
Lannoy
&
Estades
(1995)
stated
that
occasionally
bluish
discolorations
can
be
found
in
the
stipe
base.
A
collection
from
a
marshy
birch
forest
in
the
Netherlands
fitted
their
discription
well
with
respect
to
this
character.
But
the
mo-
lecular
and
microscopic
characters
placed
this
collection
in
the
current
concept
of
L.
holo-
pus.
This
normally
pale-coloured
species
usually
has
bluish
discolorations
in
the
stipe
base.
It
has
been
collected
in
the
same
locality
as
the above
mentioned
L.
pulchrum
and
ended
up
in
the
same
clade.
Apparently
the
concept
of
L.
pulchrum
by
Lannoy
&
Estades
is
merely
based
on
non-diagnostic
macroscopical
characters,
and
represents
a
mixture
of
different
species,
including
L.
scabrum,
L.
rotundifoliae
and
L.
holopus.
9.
Leccinum
schistophilum
Bon
—
Fig.
14,
Plate
8
Leccinum
schistophilum
Bon,
Doc.
Mycol.
11(44)
(1981)
35,
36.
—
Leccinum
palustre
M.
Ko-
rhonen,
Karstenia
35
(1995)
63,
Fig.
11.
Selected
icones.
Lannoy
&
Estades,
Mon.
Leccinum
(1995)
pi.
30;
Korhonen
in
Karstenia
35
(1995)
63,
Fig.
11
(as
L.
palustre).

PERSOONIA
-
Vol.
18,
Part
4,
2005556
Selected
descriptions.
Korhonen,
Karstenia
35
(1995)
63
(as
L.
palustre)
i;
Bon,
Doc.
Mycol.
11(44)
(1981)35,36.
Characteristics
—
A
slender
to
robust,
small
Leccinum,
usually
with
a
greyish
brown
pileus
and
greyish
to
blackish
stipital
squamules;
context
in
the
lower
half
of
the
stipe
with
a
bluish
discoloration.
In
wet,
often
slightly
basic
habitats.
Pileus
25-110
mm,
convex,
expanding
to
plano-convex,
with
inflexed
margin,
not
markedly
exceeding
the
tubes,
light
yellowish
brown
(Mu.
10
YR
6/4),
greyish
brown
(10
YR
5/2),
to
dark
brown
(10
YR
3-4/2),
often
evenly
coloured,
but
sometimes
with
light
spots;
minutely
tomentose,
dull.
Tubes
narrowly
to
broadly
adnate,
9-25
mm
long,
ventricose
to
broadly
ventricose,
yellowish
white
at
first,
in
later
stages
greyish
with
a
pinkish
hue. Pores
c.
0.5
mm
in
diameter,
whitish
to
yellowish
white,
discol-
ouring
brownish
when
bruised.
Stipe
46-150
mm
x
9-25
mm,
cylindrical
to
clavate,
greyish
white
to
brownish,
lightly
to
densely
covered
with
fine
squamules,
usually
not
markedly
differing
between
base
and
apex
of
stipe.
Squamules
initially
brownish,
in
later
stages
greyish
to
blackish,
sometimes
dark
brown.
Context
(greyish)
white,
when
bruised
discolouring
pinkish
(9A2)
in
the
pileus
and
upper
half
of
the
stipe,
sometimes
discolouring
bluish
green
(25A4)
in
lower
half
of
stipe;
sometimes
not
discouloring
at
all,
rarely
discolouring
greyish
after
several
hours.
Spores
(13.0-)13.5-17.0
x
5.0-6.5(-7.5)
pm,
Q
=
2.3-3.1
(-3.4),
Q
av
=
2.8-2.9
(-3.0),
subcylindrical
to
fusiform.
Basidia
25-30
x
10.0-11.5
pm,
4-spored.
Hymeno-
cystidia
30-45
x
7.5
-9.0
pm,
lageniform.
Pileipellis
cutis-like,
composed
of
cylindrical,
brownish
to
dark
brownish
elements,
(3.5-)4.5-8.0
pm
wide,
sometimes
also
with
broad
(7.0-11.5
/<m),
clearly
articulated,
elements.
Caulocystidia
30-70
x
11.0-20.5
pm,
fusiform,
clavate
or
lageniform.
Clamp-connections
absent.
Habitat
&
distribution
—
Gregarious
or
solitary,
ectomycorrhizal,
associated
with
Betula
on
mossy,
humid,
alkaline,
sandy
soils;
distribution
insufficiently
known,
prob-
ably
widespread
and
rather
rare
throughout
northern
and
western
Europe.
Collections
studied.
THE
NETHERLANDS:
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
Eiland
van
Rolvers,
2IX
1999,
H.C.
den
Bakker
30;
ibid.,23
X
1999,
H.C.
den
Bakker
61;
ibid.,4
X
2000.
H.C.
den
Bakker
121.
—
ENGLAND:
Gloucestershire,
30
VIII2000,
Alan
Hills
s.n.
—
FINLAND:
Uusimaa,Vantaa,
Veromies,
M.
Korhonen
11145
(holotype
L.
palustre,
H).
—
FRANCE:
Pas-de
Calais,
Labuissiere,
Bon
741014
(holotype
L.
schistophilum,
LIP);
Orne,
Foret
Dominial
du
Perche,
2
X
2001,
van
Brummelen
(fix
15
+
16).
—
SWEDEN:
Jamtland,
Borgsjo,
23
VIII
1999,
H.C.
den
Bakker
15;
vicinity
Are,
4IX
2000,
H.C.
den
Bakker
82.
The
typical
form
of
L.
schistophilum
can
easily
be
identified
in
the
field
and
is
charac-
terized
by
a
small,
slender
habit,
light
greyish
brown
pileus,
white
stipe
with
contrasting
fine,
greyish
to
blackish
squamules
and
a
greenish-blue
discolouring
context
in
the
lower
half
of
the
stipe.
However,
basidiocarps
with
a
brown
pileus
and
a
non-discolouring
context
may
occur,
which
accordingly
can
only
be
distinguished
from
L.
scabrum
by
its
different
ecology
(
L.
schistophilum
usually
grows
in
wet
habitats,
L.
scabrum
in
drier
habitats)
combined
with
microscopic
differences
(caulocystidia,
average
Q-value
of
spores
and
differences
in
pileipellis
structure).
Large
specimens
of
L.
schistophilum
may
be
confused
with
L.
variicolor.
Micro-
scopically
L.
schistophilum
can
easily
be
distinguished,
however,
by
the
structure
of

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
557
Fig.
14.
Leccinum
schistophilum.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

558
PERSOONIA
-
Vol.
18,
Part
4,
2005
the
pileipellis,
which
is
composed
of
long
cylindrical
elements,
while
generally
in
the
pileipellis
of
L.
variicolor
chains
of
short
cylindrical
elements
(cylindrocysts)
occur.
Furthermore
lageniform,
septate
caulocystidia
are
abundant
in
L.
variicolor,
while
these
are
only
occasionally
observed
in
L.
schistophilum.
This
taxon
was
erroneously
placed
in
subsection
Leccinum
by
Bon
(1981),
because
he
overemphasized
the
importance
of
the
grey
tinges
that
may
appear
in
the
context
several
hours
after
it
has
been
bruised.
Den
Bakker
et
al.
(2004b,
Chapter
3)
showed
that
L.
schistophilum
is
nested
in
subsection
Scabra.
10.
Leccinum
variicolor
Watling
—
Fig.
15,
Plate
9
Leccinum
variicolor
Watling,
Notes
Roy.
Bot.
Gard.
Edinburgh
29
(1969)
268.
—
I
Leccinum
variicolor
var.
bertauxii
Lannoy
&
Estades,
Doc.
Mycol.
21(81) (1991)
22.—
Leccinum
variicolor
f.
atrostellatum
Lannoy
&
Estades,
Doc.
Mycol.
21(81)
(1991)
22.
—
Leccinum
variicolor
f.
sphagnorum
Lannoy
&
Estades,
Doc.
Mycol.
21(81) (1991)
22.
Selected
icones.
Cetto,
Fungi
Vero
4
(1983)
1566
(als
L.
thalasinum);
Dahncke,
1200
Pilze
(1993)
pl.97.
Lannoy
&
Estades,
Monogr.
Leccinum
{
1995)
pi.
13-15;
R.
Phillips,
Paddest.
Schimm.
(1993)
213.
Selected
descriptions.
Watling,
Br.
Fung.
Fl.
1:
55,56.
1970.
Characteristics
—
A
small
to
medium-sized,
slender
Leccinum,
usually
with
a
char-
acteristically
variegated
pileus
and
bluish
green
discolorations
in
the
cortex
of
the
lower
half
of
the
stipe.
Pileus
35-95
mm,
convex,
expanding
to
plano-convex
or
sometimes
broadly
coni-
cal
when
mature,
with
entire
margin,
not
markedly
exceeding
the
tubes,
at
most
up
to
1
mm,
dark
brown
(Mu.
5
YR
3/1-4,10
YR
3/2-3)
with
a
radial
patttern
of
lighter
spots,
sometimes
almost
whitish
with
dark
spots
or
entirely
dark
brown;
very
finely
tomen-
tose
all
over,
often
subviscid
with
age.
Tubes
narrowly
to
broadly
adnate,
ventricose
to
broadly
ventricose,
7-18
mm
long,
greyish
or
cream-white,
discolouring
brownish
when
bruised.
Pores
c.
0.5
mm
in
diameter,
creamy
white,
often
with
yellowish
brown
spots,
discolouring
brownish
when
bruised.
Stipe
70-157
x
35
mm,
cylindrical
to
clavate,
whitish
or
greyish
white,
often
with
a
distinct
greenish
blue
discoloration
in
the
lower
half
of
the
stipe,
entirely
covered
with
brown
to
black
squamules:
fine
at
apex,
gradually
becoming
coarser
towards
base.
Context
white,
when
bruised
often
staining
pinkish
(K.
&
W.
9A3,
9A4)
in
pileus
and
upper
half
of
the
stipe,
often
discolouring
greenish
blue
(24A5,25
A5)
in
the
lower
half
of
the
stipe
(in
some
collections
limited
to
the
cortex
of
the
stipe
only);
on
drying
often
with
a
yellow
(3A7)
discoloration
in
the
lower
half
of
the
stipe.
Spores
(10.0-)
13.5
-17.5(-20.0)
x
5.0-6.5
pm,
Q
=
2.4-3.1,
Q
av
=
2.6-2.8
(-2.9),
fusiform
with
a
suprahilar
depression
or
without
a
suprahilar
depression.
Basidia
25-35
x
8.5-11.0
pm,
2-
and
4-spored.
Hymenocystidia
25-
45
x
8.0
-10.5
pm,
lageniform
to
clavate
with
a
mucronate
or
obtuse
apex.
Pileipellis
a
loose
cutis
of
cylindrical,
hyaline
or
brownish
intracellularly
pigmented
elements,
4.5-9.0(-12.5)
pm
wide,
elements
in
suprapellis
often
arranged
in
chains
and
clearly
cylindrical
(cylindrocysts),
terminal
ele-
ments
generally
conical.
Caulocystidia
(20-)
35
-
85
x
7.5
-22.0
pm,
clavate
or
irregular
cylindrical
to
lageniform
with
a
flexuose,
sometimes
forked
neck,
which
is
often
separated
from
the
body
of
the
cystidium
by
a
septum.
Clamp-connections
absent.

559
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Betula
in
mossy,
acidic
environments
or
in
Betula
swamps
with
Sphagnum,
on
peaty
and
sandy
soil;
in
the
Netherlands
moderately
common,
widespread,
but
rather
rare
in
Europe,
probably
with
a
circumboreal
distribution.
Probably
unnoticed
before
in
North
America
(see
comments).
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.
Fig.
15.
Leccinum
variicolor.

PERSOONIA
-
Vol.
18,
Part
4,
2005560
Collections
studied.
THE
NETHERLANDS:
Drenthe,
Witteveen,
Stobbeplas,
2
X
1999,
R.
Chrispijn,
H.C.
den
Bakker
45\
Erica,
Oosterse
Bos,
4
X
1999,
H.C.
den
Bakker
51;
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
Eiland
van
Rolvers,
2
IX
1999,
H.C.
den
Bakker
31;
Bergen,
Buizerdvlak,
3
XI
1978,
F.A.
van
den
Berg;
Limburg,
Mariapeel,
Griendtsveen,
8
X
1994,
M.E.
Noordeloos
94126a;
Overijsel,
Italiaanse
Meertjes,
9
X
2000,
H.C.
den
Bakker
139.
—
CANADA:
Ontario,
Manitoulin
Island,
5
X
2002,
H.C.
den
Bakker
327;
Timiskaming
District,
1IX
1981
,Malloch
1-9-81/5.
—
FIN-
LAND:
Pera-Pohjanmaa,
Rovaniemi,
M.
Korhonenl0977.
—
FRANCE:
Isere,
Massif
de
Belledonne,
Prabert,
24
X
1989,
Estades
891024/g
(holotype
L.
variicolor
var.
bertauxii,
LIP);
Massif
de
Belle-
donne,
Les
Seiglieres,
13
X
1989,
Estades
89.10.133A
(holotype
L.
variicolor
f.
atrostellatum,
LIP).
—
GREENLAND:
Sisimiut,
VIII2000,
L.
Jalink
&
AL
Nauta
8328.
—
NORWAY:
Sogn
og
Fjordane,
Sogndal,
7
IX
2000,
H.C.
den
Bakker
90.
—
SCOTLAND:
Inverness-shire,
Curr
Wood,
11
IX
2001,
H.C.
den
Bakker
226;
Mull,
Gruline
House,
7
IX
1968,
R.
Watling
6753
(holotype
L.
variicolor,
E).
—
SWEDEN:
Jamtland,
Ostavall,
22
VIII
1999,
H.C.
den
Bakker
007;
ibid.,22
VIII1999,
G.
Redeuilh,
H.C.
den
Bakker
11;
Kali,
3
IX
2000,
H.C.
den
Bakker
78.
Leccinum
variicolor
can
be
easily
recognized
in
the
field
on
account
of
its
variegated
pileus
and
distinct
blue-green
discoloration
of
the
context
of
the
stipe.
This
species
occurs
in
North
America
and
could
even
be
locally
common,
given
the
fact that
the
first
author
observed
that
this
was
one
of
the
most
common
species
of
the
subsection
Scabra
at
the
Cain
Foray
(Mycological
Society
of
Toronto)
near
Huntsville
(Ontario,
Canada)
and
on
Manitoulin
Island
(Ontario,
Canada)
in
the
fall
of
2003.
It
has
long
been
unnoticed
because
it
has
been
confused
with
L.
snellii.
Both
species
have
a
similar
discoloration
of
the
context,
and
also
the
septate
caulocystidia,
that
were
thought
to
be
diagnostic
for
L.
snellii
(Smith
et
al.,
1967)
are
found
in
L.
variicolor.
Molecu-
larly,
however,
the
two
species
cannot
even
be
considered
closely
related
(Den
Bakker
et
al.,
in
prep.).
Based
on
a
limited
number
of
herbarium
collections
and
the
original
description
of
L.
snellii,
the
most
important
differences
are
found
in
the
pileipellis,
in
particular
in
the
shape
of
the
terminal
elements.
The
pileipellis
of
L.
snellii
is
charac-
terized
by
the
presence
of
8
-
10
ptm
broad
cylindrical
elements
and
clavate
to
conical,
terminal
elements
with
dark
brown
vacuolar
pigment.
The
pileipellis
of
L.
variicolor
is
also
characterized
by
the
presence
of
short,
cylindrical
hyphal
elements,
but
usually
they
are
less
broad
(4.5
-
9.0
/on)
and
the
terminal
elements
are
conical.
In
particular
the
clavate
terminal
elements
are
distinctive
for
L.
snellii,
and
have
never
been
found
in
L.
variicolor.
There
may
also
be
an
ecological
difference
between
the
two
taxa.
In
the
Great
Lakes
region
of
Canada
the
first
author
generally
found
L.
variicolor
in
the
same
habitat
as
in
Europe,
viz.
in
humid,
acidic
habitats.
Leccinum
snellii
was
found
only
once,
and
in
a
completely
different
habitat,
viz.
a
rich,
slightly
humid
forest
on
alkaline
soil.
The
locality
of
the
specimens
on
which
the
original
description
of
L.
snellii
is
based
can
also
be
interpreted
as
a
rich
forest,
consisting
of
beech,
maple
and
yellow
birch.
More
study
is
needed
to
understand
the
morphological
delimitation
of
these
two
species.
Lannoy
&
Estades
(1995)
recognized
three
infraspecic
taxa
within
L.
variicolor,
viz.
var.
bertauxii
differing
from
var.
variicolor
by
an
evenly
coloured
blackish
pileus
and
the
absence
of
pinkish
discolorations
of
the
context
in
stipe
and
pileus,
f.
atrostel-
latum,
differing
from
the
typical
form
by
a
dark
star-shaped
pattern
on
the
pileus,
and
f.
sphagnorum,
a
form
with
uniformly
coloured
brownish
pileus.
Since
all
these
character
states
fall
within
the
normal
range
of
variability
of
L.
variicolor
examined
during
this
study,
these
taxa
are
not
considered
of
taxonomic
value.

561
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
11.
Leccinum
cyaneobasileucum
Lannoy
&
Estades
—
Fig.
16,
Plate
10
Leccinum
cyaneobasileucum
Lannoy
&
Estades,
Doc.
Mycol.
21
(81)
(1991)
23.—
Leccinum
brunneogriseolum
Lannoy
&
Estades,
Doc.
Mycol.
21
(82)
(1991)
1.
—
Leccinum
brunneogriseolum
var.
pubescentiun
Lannoy
&
Estades,
Doc.
Mycol.
21(82) (1991)
2.—
Leccinum
brunneogriseolum
f.
chlorinum
Lannoy
&
Estades,
Doc.
Mycol.
23
(89)
(1993)
65.
Selected
icones.
Courtec.
&
Duhem,
Champ.
France
Eur.
(1994)
pi.
1713;
Lannoy
&
Estades,
Monogr.
Leccinum
(1995)
pi.
18-21.
Selected
descriptions.
Lannoy
&
Estades,
Monogr.
Leccinum
(1995)
96-101.
Characteristics
—
A
medium-sized,
slender
to
robust
Leccinum
with
a
greyish
brown
pileus
and
whitish
to
greyish
stipital
squamules.
Context
usually
with
bluish
discolora-
tions
in
or
near
the
stipe
base.
Pileus
48-80
mm,
hemispherical,
expanding
to
convex
or
plano-convex,
with
in-
volute
to
deflexed
margin,
not
markedly
exceeding
the
tubes,
greyish
brown
(Mu.
10
YR
5/3)
to
light
brown
(Mu.
10
YR
6/6),
sometimes
almost
white;
surface
minutely
squamulose
tomentose,
sometimes
finely
granulose,
somewhat
viscid
when
moist.
Tubes
broadly
adnate,
12-16
mm
long,
ventricose
to
broadly
ventricose,
whitish
with
a
brownish
or
greyish
tinge.
Pores
c.
0.5
mm
in
diameter,
whitish
or
light
greyish,
dis-
colouring
brownish
when
bruised.
Stipe
72-110
x
11-23
mm,
cylindrical
to
clavate;
densely
covered
with
confluent
squamules,
which
hardly
show
the
background,
giving
the
stipe
an
almost
woolly
appearance,
squamules
first
whitish,
greyish
when
mature.
Context
whitish,
not
staining
or
discolouring
pinkish
(K.
&
W.
9A3)
in
pileus
and
apex
of
stipe
only;
staining
bluish
(23A7)
in
cortex
and
base
of
stipe,
especially
in
places
where
eaten
by
snails
or
arthropods.
Spores
(11.0-)
14.0-19.5
(-21.0)x
(3.5-)4.0-6.5(-7.0)/tm,Q
=
2.6-
4.0(-4.1),Q
av
=
3.1
-3.5,
fusiform
to
narrowly
fusiform
with
a
suprahilar
depression.
Basidia
20-30
x
11.5-13.0
pm,
4-spored,
sometimes
2-spored.
Hymenocystidia
30-45
x
5.5-7.5
pm,
lageniform
to
fusiform.
Pileipellis
a
cutis
of
easily
detachable,
wide,
cylindrical,
brown-
ish
(sepia)
elements;
terminal
elements
often
conical,
(7.0-)
8.0-15.0(-17.0)
pm
wide
and
narrowly
cylindrical,
dark
brown
elements,
4.5-6.0
pm
wide.
Caulocystidia
30-60
x
4.5-10.5
pm,
narrowly
lageniform,
utriform
or
fusiform,
when
lageniform
often
with
a
flexuose
neck.
Clamp-connections
absent.
Habitat
&
distribution
—
Gregarious
or
solitary,
ectomycorrhizal,
associated
with
Betula
on
dry
to
humid
sandy
or
peaty
soil,
sometimes
in
Sphagnum
bogs;
common
in
the
Netherlands,
moderately
common
in
large
parts
of
Europe,
although
rare
or
probably
absent
in
Finland,
Norway
and
Sweden.
Collections
studied.
THE
NETHERLANDS:
Friesland,
Schiermonnikoog,
Berkenplas,
4IX
1999,
H.C.
den
Bakker
39
;
Drenthe,,
Emmen,
Oosterbos,
2
X
1999,
H.C.
den
Bakker
43;
Gelderland,
Ede,
Gin-
kelse
zand
14IX
1993,
M.E.Noordeloos
93201;
Leusden,
Bos
Buurtweg,
22
X
2003,
J.
Wisman
s.n.;
Noord-Holland,
Amsterdamse
Waterleiding
Duinen,
Eiland
van
Rolvers,
23
X
1999,
H.C.
den
Bakker
59;
Limburg,
Mariapeel,
Griendtsveen,
Leg.
N.
Dam,
1994,
M.E.
Noordeloos
94133.
—
FRANCE:
Orne,
Foret
de
Belleme,
28IX
2001,
Van
Brummelen;
Isere,
Roybon/St
Etienne
de
St
Geoir,
1
X
1990,
Estades
9011010B
(holotype
L.
brunneogriseolum,
LIP);
Isere,
Massif
de
Belledonne,
Tourbiere
de
Luitel,
20IX
1990,
Estades
90.09.202.D.E.F.
(holotype
L.
brunneogriseolum
var.
pubescentium,
LIP);
Isere,
Col
des
Mouilles,
24
X
1989,
Estades
891024/4
(holotype
L.
brunneogriseolum
f.
chlorinum,
LIP);
Isere,
Roybon
pres
du
Col
de
toutes
Aures,
16
X
1988,
Estades
881016/1
(holotype
L.
cyaneo-
basileucum,
LIP).

562
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
cyaneobasileucum.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.
Fig.
16.

563
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Leccinum
cyaneobasileucum
is
easily
recognized
already
in
the
field
by
the
generally
dull-coloured
greyish
brown
pileus
and
woolly
stipital
surface.
Pale
and
albino
forms
occur,
which
is
confirmed
by
phylogenetic
comparison
of
ITS
and
Gapdh
sequences
of
material
of
L.
brunneogriseolum
and
L.
cyaneobasileucum.
These
pale
and
albinistic
forms
can
be
confused
with
L.
holopus
in
the
field,
but
differ
from
this
taxon
by
their
slender
spores
(Q
av
&
3.0),
and
the
abundant
cylindrocysts
in
the
pileipellis.
Due
to
the
fact that
the
name
L.
cyaneobasileucum
was
published
earlier
than
L.
brunneogriseolum,
the
first
name
has
priority
and,
unfortunately,
has
to
replace
the
latter.
Lannoy
&
Estades
(1995)
recognized
two
infraspecific
taxa
within
L.
brunneogri-
seolum,
viz.
var.
pubescentium
,
a
variety
that
differs
from
var.
brunneogriseolum
by
a
darker
brown
pileus
and
a
robust
stipe,
and
f.
chlorinum
a
form
that
shows
olivaceous
colours
in
the
margin
in
older
fruit-bodies
and
also
olivaceous,
greenish
tinges
in
the
stipe.
Darker
brown
pilei
fall
within
the
variability
of
the
species,
and
greenish
tinges
like
in
f.
chlorinum
can
also
be
observed
in
other
taxa
when
they
form
fruit-bodies
dur-
ing
a
continuous
wet
period
or
in
a
wet
habitat.
Though
a
considerable
number
of
Scandinavian
collections
of
species
of
subsection
Scabra
were
examined
and
several
field
trips
to
Sweden,
Norway
and
Finland
were
undertaken,
we never
observed
L.
cyaneobasileucum
in
Scandinavia.
12.
Leccinum
holopus
(Rostk.)
Watling
—
Fig.
17,
Plate
11
Boletus
holopus
Rostk.,
Sturm's
Deutschl.
Flora,
III
(Pilze)
5
(1844)
131;
Leccinum
holopus
(Rostk.)
Watling,
Trans.
Bot.
Soc.
Edinburgh
43
(1960)
692.
—
i
Leccinum
nucatum
Lannoy
&
Estades,
Doc.
Mycol.
23(89) (1993)
63-65.
Iconotype.
Rost.,
Sturm's
Deutschl.
Flora,
III
(Pilze)
5
(1844)
pi.
48.
Epitype.
Norway,
Sogn
og
Fjordane,
east
of
Haukaosen
Airport,
8
IX
2000,
H.C.
den
Bakker
85
(L,
designated
here).
Misapplication.
Leccinum
niveum
sensu
auct.,
Rauschert—,
Leccinum
chioneum
sensu
auct.,
Redeuilh.
—
Boletus
alhus
sensu
auct.
—
Leccinum
rotundifoliae
sensu
A.H.
Sm.,
Thiers
&
Watling,
Michigan
Bot.
6
(1967)
128;
sensu
Kallio,
Rep.
Kevo
subarct.
Res.
Stat.
12
(1975)
25-27.
Lec-
cinum
aerugineum
sensu
Lannoy
&
Estades,
Doc.
Mycol.
21(81) (1991)
23.
Selected
icones.
Korhonen,
Karstenia
35
(1995)
64:
Fig.
12;
Ryman
&
Holmasen,
Pilze
(1992)
230.
Selected
descriptions.
Watling,
Br.
Fung.
Fl.
1
(1970)
50,51.
Characteristics
—
A
small
to
medium-sized,
slender
to
robust
Leccinum;
pileus
usually
whitish
to
pale
brownish,
stipital
squamules
usually
whitish
to
light
brownish,
context
either
without
any
discolorations
or
some
bluish
spots
in
the
stipe
base.
Species
of
humid,
acidic
habitats.
Pileus
40-100
mm,
convex,
expanding
to
plano-convex,
with
entire
margin,
exceed-
ing
the
tubes
up
to
1
mm,
usually
minutely
tomentose
when
fresh,
sometimes
granular,
often
becoming
more
viscid
with
age,
whitish
to
yellowish,
greyish
white,
often
with
a
greenish
hue
or
bluish
green
in
older
fruit-bodies,
sometimes
brown
(Mu.
10
YR
5/4).
Tubes
7-18
mm
long
narrowly
to
broadly
adnate,
segmentiform
to
broadly
ventricose,
greyish
white,
finally
brownish
pinkish.
Pores
c.
0.5
mm
in
diameter,
whitish
with
yel-
lowish
brown
spots,
brownish
when
bruised.
Stipe
59-140
x
10-23
mm,
cilindrical
to
clavate,
whitish,
covered
with
coarse
whitish,
in
mature
fruit-bodies
ochre
to
greyish
squamules,
sometimes
blackish
squamules
already
present
in
young
fruit-bodies.
Con-
text
whitish,
discolouring
pinkish
in
pileus
and
stipe
when
bruised,
bluish
discolorations
often
visible
in
cortex
of
stipe
base,
but
regularly
not
discolouring
at
all.

564
PERSOONIA
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Vol.
18,
Part
4,
2005
Leccinum
holopus.
Fig.
17.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.

565
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Spores
15.5
-18.0
x
5.5-7.0
pm,
Q
=
2.5
-3.0,
Q
av
=
2.6-2.8,
broadly
fusiform
to
fusi-
form
with
a
shallow
suprahilar
depression.
Basidia
25-35
x
11.5-12.5
pm,
generally
4-spored.
Hymenocystidia40-45
x
7.5-9.0
pm,
lageniform
to
fusiform.
Caulocystidia
40-55
x
9.0-13.5
pm,
fusiform,
clavate
to
cylindrical,
often
with
a
flexuose
neck.
Pilei-
pellis
a
cutis
composed
of
cylindrical
elements,
3.5—5.0(-7.0)
/<m
in
diameter,
often
hya-
line,
granular
incrusted,
sometimes
with
brownish
vacuolar
pigment;
terminal
elements
sometimes
ellipsoid,
then
10.5-11.5
/on
in
diameter.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Betula
in
Sphagnum bogs
or
among
grass
and
mosses,
on
humid,
peaty
soil;
rather
rare
in
the
Netherlands,
common
and
probably
widespread
throughout
the
(circumboreal)
distribution
area
of
its
host.
Collections
studied.
THE
NETHERLANDS:
Zuid-Holland,
Nieuwkoop,
De
Haeck,
5
IX
1999,
M.E.
Noordeloos,
H.C.
den
Bakker
37:
Nieuwkoop,
De
Haeck,
9IX
1999,
H.C.
den
Bakker40.
—FRANCE:
Isere,
Massif
de
TObiou,
IX
1991,
Estades
9109251
(holotype
L.
nucatum,
LIP);
Isere,
Massif
de
Belledonne,
Les
Seiglieres,
Estades
8909221A
(neotype
L.
aerugineum,
proposed
by
Lannoy
&
Es-
tades,
LIP).
—
GREENLAND:
Sisimiut,
VIII2000,
E.
Jalink
&
M.
Nauta
8326.
—
NORWAY:
Sogn
og
Fjordane,
east
of
Haukaosen
Airport,
8IX
2000,
H.C.
den
Bakker
85
(epitype
L.
holopus,
L,
designated
here);
Finnmarken,
Tana
Rastiggaisa,
18
VIII
1995,
M.E.
Noordeloos
95095.
The
typical
form
of
L.
holopus
is
a
slender,
pale,
almost
whitish
bolete,
without
any
clear
discoloration
of
the
context
when
bruised,
occurring
in
Sphagnum-bogs
in
mon-
tane,
sub-boreal
and
boreal
regions.
Another,
sturdier
form
with
a
darker,
sometimes
brownish
pileus,
darker
stipital
ornamentation
and
bluish
discolorations
in
the
stipe
base,
is
found
in
wet,
acidic
birch
woods.
This
last
form
has
been
described
as
a
separate
species
by
Lannoy
&
Estades
(1993)
as
L.
nucatum.
No
molecular
evidence
to
support
the
existence
of
L.
nucatum
as a
separate
species
was
found.
Moreover,
the
differences
between
both
forms
are
not
always
clearcut
and
intermediates
occur.
For
this
reason
the
nucatum-
type
populations
do
not
justify
a
formal
taxonomic
status
and
L.
nucatum
is
placed
into
the
synonymy
of
L.
holopus.
Leccinum
holopus
can
be
distinguished
from
related
taxa
by
a
combination
of
the
following
microscopic
characters:
a
pileipellis
that
is
composed
of
slender
hyphal
ele-
ments
of
relatively
even
diameter
(3.5-5.0
pm),
abundant
cylindrical
caulocystidia
with
a
flexuose
neck
in
the
lower
half
of
the
stipe
and
an
average
Q-value
of
spores
<
2.8.
Smith
&
Thiers
(1971)
recognized
two
varieties
of
L.
holopus:
:
var.
holopus
and
var.
americanum.
According
to
Smith
&
Thiers
var.
americanum
mainly
differs
from
var.
holopus
in
the
pinkish
reddish
discoloration
of
the
context
when
bruised
and
the
blackish
stipe
ornamentation.
Both
varieties
have been
observed
in
the
field
by
the
first
author
in
Canada,
sometimes
close
together
on
the
same
spot.
Though
no
molecular
differences
could
be
found
that
were
congruent
with
these
varieties,
no
intermediates
were
observed
in
the
field.
The
absence
of
(morphological)
intermediates
seems
to
justify
recognition
of
var.
americanum
as
an
infraspecific
taxon.
SECTION
LUTEOSCABRA
Singer,
Amer.
Midi.
Naturalist
37
(1947)
122
Pileipellis
a
trichoderm
of
septate
hyphae
with
swollen
terminal
elements.
Type
species:
Leccinum
nigrescens
(Rich.
&
Roze)
Singer.

566
PERSOONIA
-
Vol.
18,
Part
4,
2005
13.
Leccinum
crocipodium
(Letell.)
Watling
—
Fig.
18,
Plate
12
Boletus
crokipodius
Letell.,
Fig.
Champ.
(1836)
pi.
166
(plate
only,
no
description);
Leccinum
crocipodium
(Letell.)
Watling,
Trans.
Bot.
Soc.
Edinburgh
39
(1961)
200.
—
Boletus
tessellatus
Gillet,
Hymenomycetes
(1878)
636,
non
B.
tessellatus
Rostk.
1844.
—
Boletus
nigrescens
Rich.
&
Roze,
Atl.
Champ.
(1888)
pi.
60,
fig.
5-10,
non
B.
nigrescens
Pallas
1771-1776;
Leccinum
nigrescens
(Rich.
&
Roze
-»)
Singer,
Am.
Midi.
Naturalist
37
(1947)
112.
—
Boletus
luteoporus
Bouchinot
in
Barbier,
Bull.
Trimestriel
Soc.
Mycol.
France
20
(1904)
91;
Leccinum
luteoporum
(Bouchinot)
Alessio,
Fungi
Europei
vol.
2A(1991)
94.
—
Boletus
velenovskyi
Smotl.,
Sitzungsber.
Konigl.
Bohm.
Ges.
Wiss.
Prag.
1911
(1912)60.
Holotype.
Letell.,
Fig.
Champ.
(1836)
pi.
166.
Epitype.
France:
Ardennes,
Sommeau/Beaumont-en-Argonne,
F.
de
Belval,
20IX
1999,
R.
Walleyn
1659
(L,
isotype
in
GENT).
Selected
icones.
Breitenb.
&
Kranzl.,
Pilze
Schweiz
3
(1991)
72.
pi.
34;
Galli,
Boleti
(1998)
260,
261;
Munoz,
Fungi
non
del.
13
(2000)
pi.
18;
Pilat
&
Dermek,
Flrivobite
Huby
(1974)
pi.
73
(as
L.
nigrescens).
Munoz,
Fungi
non
del.
13:
39-40.
2000.
Characteristics
—
Stout
species
with
yellow-brown,
often
olivaceous
tinged,
areolate-
cracked
pileus;
small,
yellow
pores
and
yellowish,
fusiform
stipe
with
fine
brown
squa-
mules;
context
staining
reddish-brown
to
violaceous
brown
then
black when
bruised;
pileipellis
a
trichoderm
of
cylindrical,
clavate
to
fusiform
elements,
7.0-15
ftm
wide;
associated
with
Quercus
and
Carpinus.
Pileus
40-75
mm
broad,
hemispherical
to
convex
with
straight,
appendiculate
margin
extending
over
tubes
for
about
1
mm,
yellowish
brown
or
reddish
brown,
sometimes
with
an
olivaceous
tinge,
later
darker
olivaceous
brown
or
blackening
when
strongly
exposed
(Mu.
2.5
-5
Y
5
-
6/4
-
6),
dry,
appressed
tomentose
becoming
entirely
cracked
with
age,
breaking
up
in
small
irregular
patches
showing
pale
grey
context
in
between.
Tubes
adnexed
to
adnate,
narrowly
to
broadly
ventricose,
up
to
15
mm
broad,
pale
yellow
(5
Y
8/6);
pores
small,
0.3-0.5
mm
in
diameter,
rounded,
rather
bright
yellow
(K.
&
W.
3A7-3B7),
turning
reddish
brown,
then
black when
bruised.
Stipe
60-150
x
15
-30
mm,
straight,
typically
fusiformwith
broadest
part
below
the
middle,
tapering
at
base,
more
rarely
cylindrical,
cream-white
at
apex,
pale
chrome-yellow
below,
minutely
squamulose
in
longitudinal
rows
of
small
yellow
to
dark
brown
floccose
squamules
which
become
coarser
towards
base.
Context
thick,
firm,
pale
yellow
or
cream-coloured,
staining
reddish brown
to
violaceous-grey
when
bruised,
blackening
in
damaged
parts.
Smell
indistinct.
Taste
mild.
Spores
(11.5
—)
12.0-15.0
x
5.0-6.5.0
pm,
on
average
13-14
x
5.5-6.0
pm,
Q
=
2.0-2.9,
Q
av
=
2.3-2.4;
fusiform with
conical
apex,
often
almost
amygdaliform,
with
slight
to
distinct
suprahilar
depression,
relatively
thin-walled,
pale
brown
in
water.
Basidia
20-35
x
5.0-9.0
pm,
clavate,
4-spored.
Hymenial
cystidia
abundant
on
edge
and
sides
of
the
pores,
20-55
x
4.5-11
x
2.0-4.0
pm,
lageniform,
often
with
rather
long,
tapering
neck,
colourless
or
with
brown,
intracellular-granular
pigment.
Pileipellis
a
rather
regular
trichoderm
of
septate
hyphae,
terminal
elements
12-50(-50)
x
6.5
-17
pm,
usually
rather
slender,
cylindrical,
clavate
or
conical,
with
intracellular,
incrust-
ing
and
extracellular
pigment.
Stipitipellis
a
cutis
of
narrow,
cylindrical,
4.0-7.0
pm
wide
hyphae,
with
small
to
fairly
large
caulohymenial
clusters
of
basidia
and
cystidia,
forming
the
squamules
on
the
stipital
surface.
Caulocystidia
25-120
x
7.0-15
x
2.0-4.5

567
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
pm,
lageniform
or
fusiform,
thin-
or
thick-walled,
colourless
or
with
brown
granular
intracellular
pigment.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Quercus
and
Carpinus
on
heavy
loamy,
often
slightly
calcareous
soil;
extremely
rare
in
the Netherlands
(Schaelsberg,
Limburg;
Sterkenburg,
Utrecht),
probably
overlooked.
Widespread,
rare
to
locally
more
common
in
Central
and
Southern
Europe,
preferably
in
thermophilous
deciduous
forest
on
heavy
loamy
soil.
Collections
studied.
BELGIUM:
Namur,
Rochefort,
Ave-et-Auffe,
14
IX
2003,
M.E.
Noordeloos
20031
7;ditto,
Boisde
Lamache,
77i.1V.
Kuyper
s.n.\
Luxembourg,
Barveaux,
13IX.
1975,
C.
Bas6666.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.
Fig.
18.
Leccinum
crocipodium.

568
PERSOONIA
-
Vol.
18,
Part
4,
2005
—
FRANCE:
Ardennes,
Sommeau/Beaumont-en-Argonne,
F.
de
Belval,
20IX
1999,
R.
Walleyn
1659\
Lorraine,
Guersling,
3IX
2000,
E.J.M.
Arnolds
00-74.
—
THE
NETHERLANDS:
Utrecht,
Sterkenburg,
3
IX
1999,
G.A.
de Vries
s.n.\
Limburg,
Valkenburg,
Schaelsberg,
6
VIII2001,
A.
Dam
01021.
The
most
distinctive
characters
of
L.
crocipodium
are
the
rather
yellow
pores,
combined
with
the
blackening
context.
It
is
the
only
species
with
this
combination
of
characters
known
from
temperate
regions
in
Europe,
where
it
has
a
preference
for
ther-
mophilous
deciduous
forests.
The
macroscopically
similar
L.
corsicum
and
L.
lepidum
have
an
exclusive
Mediterranean
distribution,
and
are
associated
with
Cistus
spp.
and
Quercus
ilex
respectively.
Leccinum
crocipodium
has
only
recently
been
discovered
in
the Netherlands
in
1999,
with
a
second
observation
in
2001.
Whether
this
may
be
due
to
climate
changes,
or
whether
it
has
been
overlooked
in
the
past,
is
difficult
to
ascertain.
Boletus
rimosus
s
28
,
often
cited
as a
synonym,
is
better
considered
a
nomen
dubium
on
account
of
the
brown-olivaceous
pileus
and
pale
pores.
See
also
Rauschert
(1987)
for
nomenclatural
comments.
14.
Leccinum
pseudoscabrum
(Kallenb.)
Šutara
—Fig.
19
Boletus
pseudoscaber
Kallenb.,
Die
Pilze
Mitteleuropas
1
(1935)
117,
pi.
40;
Leccinum
pseudo-
scabrum
(Kallenb.)
Sutara,
Ceska
Mykol.
43
(1989)
6.
—
Boletus
scaber
var.
carpini
R.
Schulz
in
Michael
&
Schulz,
Fiihr.
Pilzfreunde
1
(1924)
95;
Boletus
carpini
(R.
Schulz)
Pearson,
Naturalist
96
(1946)
818;
Leccinum
carpini
(R.
Schulz)
D.
Reid,
Trans.
Bot.
Soc.
Edinburgh
48
(1965)
525.
Excluded.
Leccinum
griseum
sensu
auct.
Holotype.
not
existent.
Type
locality:
unknown,
Germany.
Selected
icones.
Bon,
Mushr.
Toadst.
(1987)
41;
Breitenb.
&
Kranzl.,
Pilze
Schweiz
3
(1991)
70,
pi.
31;
Engel,
Rauhstielrohrlinge
(1978)
pi.
2;
Galli,
Boleti
(1998)
254,
255;
Munoz,
Fungi
non
del.
13
(2000)
pi.
19;
R.
Phillips,
Paddest.
Schimm.
(1981)
212;
Pilat
&
Dermek,
Hrivobite
Huby
(1974)
pi.
7.
Selected
descriptions
&
illustrations.
Engel,
Rauhstielrohrlinge
(1978)
14,15;
Munoz,
Fungi
non
del.
13
(2000)
42-44;
Watling,
Br.
Fungus
Fl.
(1970)
48,49,
fig.
48,54.
Characteristics
—
Slender
to
medium-sized
species
with
remarkably
dark,
rugulose
pileus
and
finely
squamulose
stipe,
associated
with
Carpinus
betulus
or
Corylus
avel-
lana
on
heavy,
clayey
soil.
Pileus
up
to
30-70
(-100)
mm
broad,
hemispherical
to
conico-convex
at
first,
expanding
to
truncately
convex
or
plano-convex,
with
involute
margin,
not
or
only
slightly
exceeding
the
tubes
for
about
1
mm,
pale
to
moderately
dark
grey-brown
(Mu.
10
YR
3-4/3,
4-5/4-6),
sometimes
with
slight
olivaceous
tinge,
dry,
dull,
initially
velvety
to
minutely
or
strongly
radially
rugulose
or
veined,
usually
cracked
with
age
with
concentric
fissures,
particularly
near
margin
showing
pale
pinkish
context.
Tubes
adnate
to
adnexed,
subventricose
to
ventricose,
up
to
25
mm
long,
yellowish
to
pink-
ish
brown
or
greyish
brown
(2.5
Y
8/4,
10
YR
8/4,
10
YR
3/3-4),
with
brownish
tinge
when
old.
Pores
very
small,
1-2
per
mm,
rounded,
pale
brownish,
staining
brown-grey
when
bruised.
Stipe
60-130
x
6-14
mm,
cylindrical,
slightly
widened
towards
base
to
subclavate,
sometimes
flexuous,
whitish
to
pale
grey
ochre,
entirely
covered
with
28)
Boletus
rimosus
Venturi,
Stud.
Micol.
(1842)
31,
pi.
10,
fig.
93,94.

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
569
brownish
black
(10
YR
6-5/4, 3-4/3,
3/2),
very
small
dot-like
squamules
arranged
in
longitudinal
rows,
which
become
somewhat
coarser
towards
the
white
tomentose
base.
Context
dirty
white
at
first
with
watery
darker
streaks,
on
cutting
first
slowly
staining
pink
to
purple
then
greyish
to
blackish
with
purple
tinge.
Spores
(12.5—)13.0—18.5
x
4.5-6.0
/rm,
on
average
14.5-15
x
4.8-5.3
pm,
Q
=
2.3-3.3,
Qav
=
2.7-2.9;
slender,
fusiform with
conical
apex,
with
distinct
suprahilar
depression,
relatively
thin-walled,
pale
brown
in
water.
Basidia
20-40
x
6.0-8.0
/rm,
clavate,
4-spored.
Hymenial
cystidia
abundant
on
edge
and
sides
of
the
pores,
18-70
x
5.0-14
x
2.0-4.0
fim,
lageniform,
often
with
rather
long,
tapering
neck,
usually
with
dark
brown,
intracellular-granular
pigment.
Pileipellis
an
intricate
trichoderm
of
septate
hyphae,
terminal
elements
15—70(—90)
x
3.5-10.5
pm,
irregularly
cylindrical
or
cla-
vate
with
intracellular
pigment.
Stipitipellis
a
cutis
of
narrow,
cylindrical,
3.5-8.0
pm
wide
hyphae,
with
small
to
fairly
large
caulohymenial
clusters
of
basidia
and
cystidia,
Fig.
19.
a.
Habitus;
b.
spores;
c.
hymenocystidia;
d.
caulocystidia.
Scale
bars
=
10
µm
resp.
1
cm.
Leccinum
pseudoscabrum.

570
PERSOONIA
-
Vol.
18,
Part
4,
2005
forming
the
squamules
on
the
stipital
surface.
Caulocystidia
25-90
x
5.0-12
x
2.0-
8.5
pm,
lageniform,
clavate
or
fusiform,
thin-
or
thick-walled,
colourless
or
with
brown
granular
intracellular
pigment.
Clamp-connections
absent.
Habitat
&
distribution
—
Solitary
or
gregarious,
ectomycorrhizal,
associated
with
Carpinus
or
Corylus
in
deciduous
forest
and
roadsides
on
calcareous
loam
and
clay
(Stellario-Carpinetum;
Alno-Padion);
rare
in
the
Netherlands,
mainly
in
F,
very
rare
in
D,
S,
K,
E
and
Z;
susceptible
(Veerman
2004);
widespread
in
Europe,
not
uncommon
in
mixed
deciduous
forests
on
clayey
or
loamy
soil.
Collections
studied.
AUSTRIA:
Karnten,
St.
Margareten
im
Rosental,
Gupf,
8
IX
1998,
M.E.
Noordeloos
9896.
—
BELGIUM:
Luxembourg,
Valley
of
Ourthe
near
Nandrin,
28
VII
1984,
L
Jalink
&
M.
Nauta
1037.
—
GERMANY:
Nordrhein-Westfalen,
Eifel,
Gerolstein,
12
VII
1967,
P.B.
Jansen
s.n.
—
THE
NETHERLANDS:
Utrecht,
Breukelen,
Gunterstein,
21
IX
1994,
M.E.
Noordeloos
9497;
—
POLAND:
Swieta
Katarzyna,
Wilkouska
Dolina,
10
IX
1966,
C.
Bas
4750.
As
on
species
level
the
epithet
pseudoscaber
is
older
than
carpini,
the
correct
name
for
the
species
is
L.
pseudoscabrum.
Note
that
there
is
no
connection
between
the
name
of
the
species
and
the
subsection
Pseudoscabra,
an
artificial
assemblage
of
species
characterized
by
cylindrocysts.
The
name
L.
griseum
has
also
been used
for
this
spe-
cies.
However,
the
basionym
Gyroporus
griseus
Quel,
is
a
superfluous
name
change
for
Boletus
scaber.
EXCLUDED
AND
DUBIOUS
NAMES
aerugineum
Boletus
chioneus
B
[var.]
aerugineus
Fr.,
Observ.
Mycol.
2
(1818)
250.;
Leccinum
aerugineum
(Fr.)
Lannoy
&
Estades,
Doc.
Mycol.
21(81) (1991)
23.
The
name
was
published
with
a
very
short
diagnosis
("pileo
stipitique
verrucoso
aeruginosis").
These
greenish
colours
are
not
diagnostic
for
certain
species,
but
can
-
depending
on
meteorological
conditions
-
occur
in
several
species.
The
name
is
therefore
best
regarded
as
nomen
dubium.
The
neotype
(from
France!)
belongs
to
L.
holopus.
chioneum
Leccinum
chioneum
(Fr.)
Redeuilh,
Doc.
Mycol.
20(78)(1990)
35;
Boletus
niveus
Fr.,
Obs.
Mycol.
1(1815)
111,
non
R.
niveus
lull,
in
Vill.,
Hist.
PI.
Dauphine
3(2)
(1789)
1040;
B.
chioneus
Fr.,
Observ.
Mycol.
2
(1818)
250
[name
change];
Leccinum
niveum
(Fr.
-»)
Rauschert,
Nova
Hedwigia
45
(1987)
503
[illegitimate,
superfluous].
This
name
has
been
considered
an
older
name
for
L.
holopus,
but the
greying
con-
text
["caro
...
tacta
cinerascifl']
contradicts
this.
Therefore
it
is
excluded
as a
nomen
dubium.
fuscoalbum
Leccinum
fuscoalbum
(Sow.)
Lannoy
&
Estades,
Doc.
Mycol.
24(94)(1994)
18;
Boletus
fuscoalbus
Sow.,
Col.
Fig.
Engl.
Fungi
3
(1814)
pi.
421.
—
Boletus
leucophaeus
Pers.,
Mycol.
Eur.
2
(1825) 140
[illegitimate,
superfluous
name
change];
Leccinum
leucophaeum
(Pers.)
Bon
in
Bon
&
Van
Haluwyn,
Doc.
Mycol.
11
(44)
(1981)
35
[illegitimate,
based
on
illegitimate,
superfluous
basionym].

571
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Ever
since Fries
(Syst.
mycol.,
Ind.
gen.,
1832),
Boletus
fuscoalbus
has
been
con-
sidered
a
synonym
of
L.
scabrum
or
as a
closely
related
species.
The
protologue
by
Sowerby
("inside
white,
sometimes
changing
to
brown,
or
even
black
on
exposure
by
cutting
in
the
air,
especially
if
in
a
tender
state
or
bruised")
makes
clear
that
the
name
is
more
likely
an
older
name
for
L.
pseudoscabrum
or
L.
duriusculum.
Since
there
are
apparently
four
different
interpretations
for
Boletus
fuscoalbus/B.
leucophaeus
(a
synonym
of
L.
scabrum;
a
closely
related
species
to
L.
scabrum;
L.
pseudoscabrum;
L.
duriusculum),
it
seems
best
to
consider
the
name
as a
nomert
dubium.
molle
Leccinum
molle
(Bon)
Bon,
Doc.
Mycol.
19
(75)
(1989)
58;
Krombholziella
mollis
Bon,
Doc.
Mycol.
14
(56)
(1984)
22.
The
description
of
Bon
(1984)
and
Lannoy
&
Estades
(1995)
is
applicable
to
a
number
of
species
since
the
greenish
tinges
and
discolorations
represent
a
character
complex
that
is
commonly
encountered
in
several
species
of
subsection
Scabra
when
the
fruit-bod-
ies
have
developed
during
a
cool,
humid
period.
The
type
material
is
in
poor
condition
and
not
sufficiently
well-preserved
to
study
its
molecular
and
microscopical
characters.
The
spores
could
be
measured
and
have
an
average
Q-value
of
3.1,
which
is
typical
for
L.
scabrum.
It
is
excluded
as
a
nomen
dubium.
murinaceum
Leccinum
murinaceum
(J.
Blum)
Bon,
Doc.
Mycol.
9
(35)
(1979)
41;
Boletus
murinaceus
Blum,
Bull.
Trimestriel
Soc.
Mycol.
France
85
('1969'
1970)
560.
The
holotype
of
L.
murinaceum
(Blum
750,
PC)
is
in
very
poor
condition,
and
imma-
ture,
and
it
proved
to
be
impossible
to
study
any
molecular
or
morphological
characters.
The
original
diagnosis
of
Blum
(1970)
does
not
offer
much
help
to
clarify
the
identity
of
this
taxon
either.
The
interpretation
of
Lannoy
&
Estades
(1995:
68)
comes
close
to
the
current
concept
of
L. melaneum.
According
to
these
authors
L.
murinaceum
is
not
only
mycorrhizal
with
Betula,
but
also
with
Salix,
a
mycorrhizal
association
that
is
exceptional
in
subsection
Scabra.
It
is
considered
a
nomen
dubium
for
these
reasons.
olivaceosum
Leccinum
olivaceosum
Lannoy
&
Estades,
Doc.
Mycol.
24
(94)
(1994)
10.
The
type
material
is
in
poor condition
and
could
not
be used
for
further
study.
The
plates
of
Lannoy
&
Estades
(1995)
show
a
species
that
looks
macro-morphologically
like
L.
holopus
and
L.
schistophilum.
A
molecular
study
by
Binder
(1999)
based
on
material
provided
by
the
authors
shows
that
its
ribosomal
sequences
are
reminiscent
of
those
of
L.
holopus.
Further
data
are
needed
to
demonstrate
if
this
taxon
is
really
synonymous
with
L.
holopus.
oxydahile
Leccinum
oxydahile
(Singer)
Singer,
Amer.
Midi.
Naturalist
37
(1947)
123;
Krombholzia
oxydabilis
Singer,
Schweiz.
Z.
Pilzk.
16
(1938)
149.
The
original
description
L.
oxydahile
is
equally
applicable
to
L.
holopus,
L.
variicolor,
L.
cyaneobasileucum,
L.
snellii
and
L.
schistophilum.
Unfortunately,
no
type
material
was
designated
by
Singer.
According
to
Hahn
(1997)
L.
oxydabile
is
a
taxon
with
a

572
PERSOONIA
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Vol.
18,
Part
4,
2005
brownish
pileus,
a
stipe
base
that sometimes
shows
traces
of
blue,
and
which
has
broad
(up
to
20
pm)
pileipellis
elements.
This
decription
comes
close
to
L.
cyaneobasileucum.
Hahn
described
the
stipe,
however,
as
greyish
brown
to
blackish,
which
is
atypical
for
L.
cyaneobasileucum.
Another
option
would
be
that
L.
oxydabile
sensu
Hahn
represents
L.
schistophilum,
butthe
broad
pileipellis
elements
do
not
fit
with
our
interpretation
of
that
species.
Given
the
confusion
associated
with
the
name
L.
oxydabile
(Hahn,
1996),
we
think
it
is
better
to
abandon
the
name
and
consider
it
a
nomen
confusum.
rufum
Boletus
rufus
Schaeff.
Fungorum
qui
in
Bavaria
et
Palatinatu
circa
Ratisbonam
nacuntur
icones
nativis
coloribus
expressae
(1774)
Vol.
IV;
Leccinum
rufum
(Schaeff.)
Kreisel,
Boletus,
SchrReihe
1
(1984)30.
The
name
Leccinum
rufum
has
often
beenused
for
L.
aurantiacum.
Dorfelt
&
Berg
(1990)
interpreted
this
name
as
the
taxon
we
call
L.
versipelle.
Schaeffer's
original
plate
(Schaeffer,
1763)
and
description
(Schaeffer,
1774)
of
Boletus
rufus
can
be
applied
to
our
current
concept
of
L.
vulpinum,
L.
versipelle
and
L.
aurantiacum.
Further
use
of
the
name
Leccinum
rufum
would
only
cause
more
nomenclatural
confusion
and
as
such
the
name
should
be
rejected.
thalassinum
Leccinum
thalassinum
Pilat
&
Dermek,
Hribovite
huby
(1974)
146
Lannoy
&
Estades
(1995)
listed
this
taxon
as
a
possible
synonym
of
L.
variicolor.
However,
it
cannot
be
excluded
that
it
may
represent
L.
schistophilum.
Unfortunately
the
type
is
in
poor
condition,
and
the
shape
of
the
caulocystidia,
essential
to
distinguish
L.
schistophilum
from
L.
variicolor,
could
not
be
observed.
ACKNOWLEDGEMENTS
We
would
like
to
thank
Thom
Kuyper
for
his
advice
and
help
with
the
nomenclature
and
improve-
ment
of
the
text.
Marijke
Nauta
is
thanked
for
her
editorial
advice
and
critical,
but
helpful
remarks
on
the
text.
We
would
also
like
to
thank
Jan
Frits
Veldkamp
for
his
help
with
the
Latin
diagnosis.
Further
we
are
most
grateful
to
Joanne
Porck
for
her
beautiful
illustrations.
Jaap
Wisman,
Ruben
Walleyn,
Alan
Hills
and
the
curators
of
LIP,
SFSU,
MICH,
H,
PC
are
thanked
for
providing
(type)material.
We
are
grateful
to
Nancy
Ironside
for
giving
the
first
author
the
opportunity
to
collect
in
Ontario,
Canada.
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D.K.
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Th.W.
Kuyper.
2004.
A
comparison
of
the
application
of
a
biological
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morphological
species
concept
in
the
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crustuliniforme
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a
phylogenetic
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Persoonia
18:
285-316.
Bessette,
A.R.,
W.C.
Roody
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A.E.
Bessette.
2000.
North
American
Boletes:
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the
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Pored
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University
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Syracuse.
Binder,
M.
1999.
Zur
molekularen
Systematik
der
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Boletineae
und
Sclerodermatineae
subordo
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[PhD
Dissertation],
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University
of
Regensburg.
Binder,
M.
&
H.
Besl.
2000.28S
rDNA
sequence
data
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2000:
75-86.
Grafica
Sette,
Brescia.

573
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
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M.
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J.
1967a.
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determination
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135-161.
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575
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Appendix
Species
GenBank
Accession
#
Species
GenBank
Accession
#
Boletellus
betula
Boletellus
russellii
Boletus
aereus
Boletus
appendiculatus
Boletus
coniferarum
Boletus
edulis
Boletus
erythropus
Boletus
eximius
Boletus
flavoniger
Boletus
griseus
Boletus
inedulis
Boletus
lurides
Boletus
mirabilis
Boletus
ornatipes
Boletus
radicans
Boletus reticulatus
Boletus
retipes
Boletus
satanas
Boletus
seperans
Boletus
subglabripes
Boletus
subvelutipes
Boletus
viridiflavus
Chalciporus
amarellus
Chalciporus
piperatus
Chamonixia
caespitosa
Heimiella
retisporus
Leccinum
albellum
Leccinum
aurantiacum
s.s.
(L.
populinum)
Leccinum
carpini
Leccinum
chromapes
Leccinum
chromapes
Leccinum
corsicum
Leccinum
crocipodium
Leccinum
duriusculum
1
Leccinum
duriusculum
2
Leccinum
duriusculum
3
Leccinum
flavostipitatum
Leccinum
holopus
Leccinum
holopus
(L.
nucatum)
Leccinum
lepidum
Leccinum
albostipitatum
(L.
aurantiacum)
Leccinum
longicurvipes
AF050642
AF050651
AF456836
AF456837
AF456827
AF456816
AF139683
AF
139684
AF456829
AF456834
AY612803
AF
139686
AF335451
AF456817
AF336241
AF456824
AF456823
AF071528
AF457404
AF139688
AY612804
AY612805
Af456835
AF336244
AF336245
AF050650
AY
612811
AF
139703
AF
139691
AY612834
AF139709
AF139693
AF
139694
AF
139690
AF139695
AF
139699
AF139696
AF139697
AF139700
AF139698
AF139689
AF139685
Leccinum
schistophilum
(palustre)
Leccinum
versipelle
(percandidum)
Leccinum
rotundifoliae
Leccinum
rubropunctus
1
Leccinum
rubropunctus
2
Leccinum
rugosiceps
Leccinum
variicolor
Leccinum
versipelle
(cerinum)
Leccinum
scabrum
Phyloporus
pelletieri
Rubinoboletus
ballouii
Tylopilus
alboater
Tylopilus
badiceps
Tylopilus
felleus
Tylopilus
ferrugineus
Tylopilus
nigerrimus
Tylopilus
plumbeoviolaceus
Tylopilus
rhoadsiae
Tylopilus
rufonigricans
Tylopilus
tabacinus
Xanthoconium
affine
1
Xanthoconium
affine
2
Xerocomus
amazonicus
Xerocomus
badius
Xerocomus
chrysentron
Xerocomus
cisalpinus
Xerocomus
depilatus
Xerocomus
dryophilus
Xerocomus
fennicus
Xerocomus
hortonii
Xerocomus
illudens
Xerocomus
impolitus
Xerocomus
parasiticus
Xerocomus
porosporus
Xerocomus
pruinatus
Xerocomus
pruinatus
Xerocomus
ripariellus
Xerocomus
ripariellus
Xerocomus
rubellus
Xerocomus
rubellus
Xerocomus
rubellus
Xerocomus
subtomentosus
AF139701
AF139702
AF139704
AF139687
AY612812
AY612813
AF
139706
AF139692
AF139705
AF456818
AY612823
AF139708
AY612833
AF139710
AF
139711
AF456832
AF457405
AY612836
AY612835
AY612837
AY612838
AF457406
AY612839
AF050648
AF514808
AF514812
AF139712
AF514823
AF514821
AF139713
AF139714
AF139715
AF050646
AF050645
AF402140
AF514825
AF514816
AF050649
AF514828
AF514829
AF514830
AF139716
28S
Accessions
used
for
phylogenetic
reconstructions.
When
synonyms
were
used
for
the
original
Genbank
accessions,
these
are
put
between
brackets
behind
the
epithet.
Species
GenBank
Accession
#
Species
GenBank
Accession
#
Boletellus
betula
AF050642
Leccinum
schistophilum
(palustre)
AF139701
Boletellus
russellii
AF050651
Leccinum
versipelle
(percandidum)
AF139702
Boletus
aereus
AF456836
Leccinum
rotundifoliae
AF139704
Boletus
appendiculatus
AF456837
Leccinum
rubropunctus
1
AF
139687
Boletus
coniferarum
AF456827
Leccinum
rubropunctus
2
AY612812
Boletus
edulis
AF456816 Leccinum
rugosiceps
AY612813
Boletus
erythropus
AF139683
Leccinum
variicolor
AF
139706
Boletus
eximius
AF139684
Leccinum
versipelle(cerinum)
AF
139692
Boletus
flavoniger
AF456829
Leccinum
scabrum
AF139705
Boletus
griseus
AF456834
Phyloporus
pelletieri
AF456818
Boletus
inedulis
AY612803
Rubinoboletus
ballouii
AY612823
Boletus
lurides
AF
139686
Tylopilus
alboater
AF139708
Boletus
mirabilis
AF335451
Tylopilus
badiceps
AY612833
Boletus
ornatipes
AF456817
Tylopilus
felleus
AF139710
Boletus
radicans
AF336241
Tylopilus
ferrugineus
AF139711
Boletus reticulatus
AF456824
Tylopilus
nigerrimus
AF456832
Boletus
retipes
AF456823
Tylopilus
plumbeoviolaceus
AF457405
Boletus
satanas
AF071528
Tylopilus
rhoadsiae
AY612836
Boletus
seperans
AF457404
Tylopilus
rufonigricans
AY612835
Boletus
subglabripes
AF139688
Tylopilus
tabacinus
AY612837
Boletus
subvelutipes
AY612804
Xanthoconium
affine
1
AY612838
Boletus
viridiftavus
AY612805
Xanthoconium
affine
2
AF457406
Chalciporus
amarellus
Af456835
Xerocomus
amazonicus
AY612839
Chalciporus
piperatus
AF336244
Xerocomus
badius
AF050648
Chamonixia
caespitosa
AF336245
Xerocomus
chrysentron
AF514808
Heimiella
retisporus
AF050650
Xerocomus
cisalpinus
AF514812
Leccinum
albellum
AY
612811
Xerocomus
depilatus
AF139712
Leccinum
aurantiacum
s.s.
AF
139703
Xerocomus
dryophilus
AF514823
(L.
populinum)
Xerocomus
fennicus
AF514821
Leccinum
carpini
AF
139691
Xerocomus
hortonii
AF139713
Leccinum
chromapes
AY612834
Xerocomus
illudens
AF139714
Leccinum
chromapes
AF
139709
Xerocomus
impolitus
AF139715
Leccinum
corsicum
AF139693
Xerocomus
parasiticus
AF050646
Leccinum
crocipodium
AF139694
Xerocomusporosporus
AF050645
Leccinum
duriusculum
1
AF
139690
Xerocomus
pruinatus
AF402140
Leccinum
duriusculum
2
AF139695
Xerocomus
pruinatus
AF514825
Leccinum
duriusculum
3
AF
139699
Xerocomus
ripariellus
AF514816
Leccinum
flavostipitatum
AF139696
Xerocomus
ripariellus
AF050649
Leccinum
holopus
AF139697
Xerocomus
rubellus
AF514828
Leccinum
holopus
AF139700
Xerocomus
rubellus
AF514829
(L.
nucatum)
Xerocomus
rubellus
AF514830
Leccinum
lepidum
AF139698
Xerocomus
subtomentosus
AF139716
Leccinum
albostipitatum
AF139689
(L.
aurantiacum)
Leccinum
longicurvipes
AF139685

576/1
PERSOONIA
-
Vol.
18,
Part
4,
2005
(Austria,
M.
E.
Noordeloos).
Quercus
Populus
(The
Netherlands,
M.
E.
Noordeloos);
c.
fruit-body
associated
with
Leccinum
aurantiacum.
Plate
1.
a.
Fruit-body
with
aberrant
light
stipe
(Finland,
M.
E.
Noordeloos);
b.
fruit-body
associated
with

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
577/2
Plate
2.
a.
(Austria,
K.
F.
Reinwald);
b.
Leccinum
albostipitatum
L.
vulpinum
(Finland,
M.
E.
Noor-
deloos).

PERSOONIA
-
Vol.
18,
Part
4,
2005
578/3
Leccinum
versipelle.
Plate
3.
a-d.
Forms
with
orange
to
brownish
caps;
e
and
f.
light
forms
(Finland
all
photos
M.
E.
Noordeloos).

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
579/4
Leccinum
duriusculum.
a.
Robust
fruit-bodies
(Italy,
M.
E.
Noordeloos);
b.
slender
fruit-bodies
(The
Netherlands,
M.
E.
Noordeloos).
Plate
4.

PERSOONIA
-
Vol.
18,
Part
4,
2005
580/5
Plate
5.
Leccinum
scabrum.
a.
Habitus
(Ontario,
Canada,
H. C.
den
Bakker);
b,
extreme
discoloration
of
the
context
(Finland,
M.
E.
Noordeloos);
c.
habitus
(The
Netherlands,
C.
Bas).

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
581/6
Leccinum
melaneum.
Plate
6.
a.
&
b.
Scotland,
H. C.
den
Bakker;
c.
The
Netherlands,
M.
E.
Noor-
deloos.

582/7
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
rotundifoliae.
a
&
c.
Greenland,
L.
Jalink
&
M.
Nauta;
b.
Finland,
M.
E.
Noor
deloos.
Plate
7.

583/8
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
Leccinum
schistophilum.
Plate
8.
a.
Typical
habit
(Sweden,
H. C.
den
Bakker);
b.
typical
habit
(Finland,
M.
E.
Noordeloos);
c.
atypical
habit
(The
Netherlands,
H. C.
den
Bakker).

584/9
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
variicolor.
Plate
9.
a.
Pileus
(The
Netherlands,
M.
E.
Noordeloos);
b.
typical
discoloration
of
context
(Ontario,
Canada,
H. C.
den
Bakker);
c.
(The
Netherlands,
M.
E.
Noordeloos);
d.
(Slovakia,
M.
E.
Noordeloos).

Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
585/10
Plate
10.
Leccinum
cyaneobasileucum.
a.
Normal
form
(The
Netherlands,
M.
E.
Noordeloos);
b.
rare
white
(albinistic)
form
(The Netherlands,
M.
E.
Noordeloos);
c.
normal
form
(Belgium,
R.
Walleyn).

586/11
PERSOONIA
-
Vol.
18,
Part
4,
2005
Leccinum
holopus.
a-f.
Sturdy
brownish
forms;
g.
typical
slender,
whitish
form;
c
&
d
(extreme)
discoloration
of
the
context
(a
&
e,
The
Netherlands,
M.
E.
Noordeloos,
b,
c,
d,
Scotland,
H. C.
den
Bakker,
f,
Sweden,
H.
C.
den
Bakker,
g,
Sweden,
M.
E.
Noordeloos).
Plate
11.

587/12
Den
Bakker
&
Noordeloos:
A
revision
of
European
species
of
Leccinum
(Austria,
K.
F.
Reinwald).
L.
pseudoscabrum
Leccinum
crocipodium
Plate
12.
a.
(pileal
margin
of
mature
fruit-body
is
abnormally
ruptured!)
(The
Netherlands.
Peter
Klok);
b.